LIBRARY 

UNIVERSITY  OF  CALIFOKNTAI 


Digitized  by  the  Internet  Archive 

in  2007  with  funding  from 

Microsoft  Corporation 


http://www.archive.org/details/coalmeasuresamphOOmoodrich 


THE  COAL  MEASURES  AMPHIBIA 

OF 

NORTH  AMERICA 


By 

ROY  LEE  MOODIE 

Associate  in  Anatomy,  University  oj  Illinois,  Chicago 


Published  by  the  Carnegie  Institution  of  Washington 
WASHINGTON,  1916 


LIBRARY 
UNIVERSITY  OF  CAL 
DAVIS 


CAKNEGIE  INSTITUTION  OF  WASHINGTON 
Publication  No.  238 


Copies  of  this  Book 

were  first  issued 

SEP  28  1916 


PRESS  OF  J.  B.  LIPPINCOTT  COMPANY 
PHILADELPHIA 


PREFATORY  NOTE. 

The  Carnegie  Institution  of  Washington  has  already  published  several  mono- 
graphs upon  paleobiological  subjects,  written  by  its  research  associates,  Hay, 
Wieland,  and  Case.  Each  author  has  dealt  with  the  subject-matter  of  his  particular 
field,  but  each  has  brought  to  bear  upon  his  work  common  factors  which  have 
placed  his  labors  upon  a  broader  basis  than  the  mere  morphological  descriptions 
of  fossil  forms  of  life.  Case  has  published  four  monographs  upon  the  morphology 
and  taxonomy  of  the  Permo-Carboniferous  vertebrates  of  North  America,  and 
has  followed  these  by  a  fifth,  in  which  all  the  known  factors  bearing  upon  the 
development  of  the  life  were  assembled  in  an  effort  to  discuss  the  paleogeography 
of  the  period.  In  his  conception  paleogeography  is  a  very  broad  term,  involving 
not  only  a  study  of  the  distribution  of  land,  water,  and  life  in  any  one  interval  of 
time,  but  a  consideration  of  all  the  factors  in  the  extremely  complex  inter-rela- 
tions of  organic  and  inorganic  matter  and  causes  which  influence  the  development 
of  each  part. 

Geologists  and  paleobiologists  have  alike  suffered  in  their  interpretation  of 
past  conditions,  because  of  their  lack  of  knowledge  of  the  work  done  by  others. 
Stratigraphy  may  not  be  interpreted  from  the  preserved  fossils  without  a  knowledge 
of  biological  laws,  and  the  formations  of  the  earth  may  not  safely  be  rearranged 
to  account  for  the  present  or  past  distribution  of  life  without  a  knowledge  of  geo- 
logical processes. 

It  is  obvious  that  such  work  is  beyond  the  possibilities  of  any  one  man;  it  is 
rather  the  work  of  a  group  of  men,  each  broadly  trained  and  each  master  of  his 
own  field  and  able  to  contribute  to  and  criticize  the  work  of  his  fellows.  Nowhere 
could  close  cooperation  of  this  kind  be  better  accomplished  than  under  a  system 
such  as  the  Carnegie  Institution  of  Washington  has  developed,  whereby  the  research 
associates  of  the  Institution  and  others  of  its  staff  may  call  in  the  assistance  of 
men  in  related  fields.  Already  the  value  of  this  procedure  is  apparent  in  the  results 
accomplished  by  cooperation. 

The  following  monograph,  by  Dr.  Roy  L.  Moodie,  adds  an  important  link  to 
the  series  of  paleobiological  publications  of  the  Institution  and  is  closely  connected 
with  the  work  already  done  upon  the  Permo-Carboniferous  vertebrates,  since  it 
supplies  a  description  of  the  life  of  the  period  immediately  preceding.  It  is  hoped 
that  the  volume  will  contribute  in  no  small  measure  to  an  understanding  of  the 
broader  problems  of  paleogeography  and  the  recognition  of  the  mutual  problems 
of  the  paleobiologists  and  the  geologists. 

E.  C.  Case. 
University  of  Michigan,  March  i$,  igi6. 


PREFACE. 

The  question  of  the  origin  of  land  vertebrates,  which  has  appealed  so  strongly 
to  students  of  fossil  Amphibia,  is  by  no  means  solved  from  the  material  furnished 
by  the  Coal  Measures  of  North  America.  The  Amphibia  are,  however,  well  known 
from  several  localities  in  the  Coal  Measures  of  this  continent,  where  skeletons  have 
been  recovered  which  are  sufficiently  well  preserved  to  afford  a  fair  knowledge  of 
their  anatomy.  The  specimens  rescued  from  the  dumps  of  the  coal  mines  are  regret- 
tably few  in  comparison  with  the  number  that  must  have  been  burned  as  fuel,  or 
carried  down  the  slopes  as  silt.  Yet  scanty  as  is  the  material  thus  collected,  it  is  of 
great  importance,  because  it  represents  such  an  early  period  in  the  recorded  history 
of  the  air-breathing  vertebrates. 

The  amphibian  fauna  in  the  Coal  Measures  of  North  America  is  represented  by 
several  hundred  individual  specimens,  preserved  in  various  museums.  All  of  the 
collections  have  been  available  in  the  preparation  of  this  memoir,  with  the  exception 
of  those  species  from  Nova  Scotia  which  are  preserved  in  the  Peter  Redpath 
Museum  of  McGill  University  and  in  the  British  Museum  of  Natural  History.  The 
European  material,  which  has  been  used  in  comparisons  with  the  American  forms, 
has  been  studied  chiefly  from  the  literature,  although  there  have  been  available  a 
series  of  specimens  of  Branchiosaurus  amblystomus  Credner,  from  Saxony,  presented 
by  the  late  Professor  Credner,  and  a  single  specimen  of  Archegosaurus  from  Dr.  von 
Huene,  of  Tubingen. 

The  collection  which  has  been  of  the  greatest  value  is  that  at  the  American 
Museum  of  Natural  History,  chiefly  assembled  by  Dr.  J.  S.  Newberry  from  the 
dumps  of  the  coal  mines  at  Linton,  Ohio,  while  he  was  in  charge  of  the  Ohio  Geo- 
logical Survey  (1869- 1884).  This  collection,  a  part  of  which  is  at  Columbia  Uni- 
versity, furnished  Cope  with  the  most  of  his  type  material  for  the  "Synopsis  of 
the  Extinct  Batrachia  from  the  Coal  Measures"  (123).*  This  entire  collection, 
including  all  of  Professor  Cope's  types  and  representing  many  new  and  hitherto 
undescribed  forms,  was  generously  placed  at  the  writer's  disposal  for  a  period  of 
five  years  through  the  kindness  of  Dr.  Bashford  Dean  and  Dr.  Louis  Hussakof.  Dr. 
Hussakof  made  a  trip  through  the  Linton  region  and  his  description  of  the  place 
occupied  by  the  "Old  Diamond  Mine"  is  given  on  page  16. 

An  interesting  collection  of  air-breathing  vertebrates  from  the  Coal  Measures, 
representing  19  species,  is  in  the  U.  S.  National  Museum  (464).  This  is  chiefly  the 
collection  of  Mr.  R.  D.  Lacoe  and  includes  specimens  from  Mazon  Creek,  Illinois, 
from  Kansas,  and  from  Linton,  Ohio.  It  is  especially  important  in  that  it  contains 
the  skeleton  (plate  20,  fig.  3)  of  the  oldest  known  reptile,  Eosauravus  copei  Williston 
(Jour.  Geol.,  xvi,  295).  It  contains  also,  besides  many  of  Cope's  types,  new  forms 
which  have  been  described  by  the  writer  (464,  470,  471,  472,  473,  474,  478,  479). 
Dr.  Stuart  Weller  first  secured  the  use  of  this  collection  for  me,  and  its  continued  use 
has  been  granted  by  Dr.  C.  D.  Walcott.  Mr.  Charles  Gilmore  has  called  my  atten- 
tion to  several  interesting  specimens  and  has  kindly  loaned  them  for  description. 

»  The  numbers  in  parentheses  refer  to  the  bibliography  at  the  end  of  this  volume. 

V 


VI  PREFACE. 

A  small  but  interesting  collection  of  Mazon  Creek  Amphibia  is  that  of  the  Pea- 
body  Museum  of  Yale  University.  Through  the  courtesy  of  the  officers  of  this 
museum  the  writer  was  permitted  to  study  these  specimens  and  was  given  a  grant 
for  their  illustration.  The  results  of  that  study  are  contained  in  a  previous  paper 
(478)  and  in  the  present  memoir.  Dr.  Schuchert  has  offered  suggestions  as  to  the 
environmental  conditions  of  the  ancient  Amphibia. 

A  few  specimens  of  Coal  Measures  Amphibia  are  at  the  Walker  Museum,  Uni- 
versity of  Chicago.  This  collection  includes  the  type  of  Micrerpeton  caudatum 
Moodie,  the  first  branchiosaur  discovered  in  the  western  hemisphere,  and  a  few 
specimens  from  Linton,  Ohio. 

A  single  specimen  of  Amphibamus  grandiceps  Cope,  very  beautifully  preserved, 
is  in  the  possession  of  Mr.  L.  E.  Daniel's,  of  Rolling  Prairie,  Indiana.  This  specimen 
has  been  studied  and  described  by  Hay  (316)  and  by  the  writer  (462,  469,  478). 

The  works  of  Cope  and  Dawson,  published  between  i860  and  1897,  on  the 
Amphibia  from  the  Coal  Measures,  have  been  indispensable  in  the  present  study. 
It  has  been  necessary  to  rely  on  the  published  descriptions  and  photographs  of  the 
interesting  fauna  from  Nova  Scotia,  since  it  has  not  been  possible  for  me  to  visit 
and  examine  the  types  preserved  in  the  Peter  Redpath  Museum  of  McGill  Uni- 
versity and  in  the  British  Museum  of  Natural  History.  It  has  been  possible  to 
check  Dawson's  work,  to  a  certain  extent,  by  a  study  of  a  series  of  excellent  photo- 
graphs of  the  types  of  Coal  Measures  Amphibia  collected  by  Dawson  and  Lyell 
and  described  by  Dawson  and  Owen.  The  descriptions  of  these  authors  have  been 
drawn  on  for  the  discussion  of  the  Canadian  forms. 

The  descriptions  given  below  have  been  made  full  and  complete  in  the  belief 
that  in  this  way  our  knowledge  of  these  interesting  vertebrates  may  be  advanced. 
Many  of  the  species  have  been  described  elsewhere  in  scattered  papers  by  various 
authors.  These  descriptions  have  been  revised  and  verified  and  are  collected  here 
in  monographic  form.  The  work  is  a  morphologic  and  taxonomic  revision  of  the 
Amphibia  from  the  Coal  Measures  of  North  America.  Especial  attention  has  been 
paid  to  the  factors  which  have  been  most  active  in  the  evolution  of  the  group,  so 
far  as  these  factors  may  be  interpreted.  It  is  the  author's  hope  that  this  review  may 
open  up  the  field  for  many  more  workers,  since  we  are  just  beginning  to  learn  about 
the  evolution  of  this  group  of  vertebrates. 

The  trustees  of  the  Elizabeth  Thompson  Science  Fund  allotted  a  grant  for  the 
present  investigation.  This  aid  has  enabled  the  writer  to  present  his  work  in  much 
better  form  than  would  have  been  possible  otherwise.  Dr.  S.  W.  Williston  has 
offered  many  suggestions  and  criticisms  which  have  been  gratefully  adopted.  It 
is  with  the  greatest  sense  of  pleasure  that  the  author  dedicates  this  memoir  to  his 
teacher  and  friend.  After  the  manuscript  was  completed  the  author  enjoyed  a  visit 
from  Mr.  D.  M.  S.  Watson,  of  King's  College,  London,  whose  knowledge  of  the 
European  and  African  forms  enabled  him  to  offer  several  very  valuble  suggestions. 

It  is  fitting  also  to  express  my  indebtedness  to  the  Carnegie  Institution  of 
Washington  for  the  privilege  of  publishing  my  work  in  the  series  of  monographs 
contributed  by  Dr.  E.  C.  Case,  dealing  with  the  anatomy  and  relationships  of  the 
early  land  vertebrates  of  North  America. 

Roy  Lee  Moodie. 


Prefatory   Note 
Preface 


9-22 
23-36 


CONTENTS. 

Page. 

iii 

\    vi 

List  of  Illustrations viii-x 

CHAP. 

I.  The  Prohi.em  of  the  Amphibia  from  the  Coal  Measures ,-e 

II.  History  of  the  Discovery  of  Amphibia  in  the  Coal  Measures 6-8 

III.  Stratigraphic  and  Geographic  Distribution  of  Amphibia  in  the  Coal  Measures  of  North 

America 

IV.  The  Morphology  of  the  Coal  Measures  Amphibia 

V.  The  Amphibia  of  the  Devonian  and  Mississippian  of  North  America 37-38 

VI.  A  History  of  the  Classification  of  the  Amphibia,  with  Especial  Reference  to  the  Species 

from  the  Coal  Measures 39-45 

VII.  Classification  of  Amphibia  Adopted  in  this  Work,  and  a  List  of  the  Coal  Measures 

Amphibia  from  North  America 46-48 

VIII.  Definition  of  the  Class  Amphibia,  the  Subclass  Euamphibia,  and  the  Order  Branchio- 

sauria 49-50 

IX.  The  American  Coal  Measures  Branchiosaurid/E 51-66 

X.  The  Order  Caudata 67-71 

XI.  The  Order  Salientia 72-74 

XII.  The  Subclass  Lepospondylia,  the  Order  Microsauria,  and  the  Group  Aistopoda 75-77 

XIII.  The  Microsaurian  Family  Hylonomid*,  from  the  Coal  Measures  of  Nova  Scotia 78-84 

XIV.  The  Microsaurian  Family  Tuditanid;e,  from  the  Coal  Measures  of  Ohio  and  Pennsylvania  85  in 
XV.  The  Mh  kosaurian  Family  Stegopid/E,  from  the  Coal  Measures  of  Ohio 112-114 

XVI.  The  Microsaurian  Family   Urocordylid.e,  from  the  Coal  Measures  of  Ohio 115  125 

XVI  I.  The  Microsaurian  Family  Amphibamid.e,  from  the  Coal  Measures  of  Ma/on  Creek,  Illinois  126-134 

XVIII.  The  Microsaurian  Family  Nyraniid^,  from  the  Coal  Measures  of  Ohio 135-138 

XIX.  The  Aistopodous  Microsaurian  Family  Ptyoniid.e,  from  the  Coal  Measures  of  Ohio.  .    .  139-146 
XX.  The  Microsaurian  Family  Molgophid^e,  from  the  Coal  Measures  of  Ohio  and  Mazon 

Crbbk,  Illinois 147  154 

XXI.  The  Microsaurian  Family  Sauropleurid/E,  from  the  Coal  Measures  of  Ohio 155-170 

XXII.  The  Microsaurian  Family  Ichthycanthid.e,  from  the  Coal  Measures  of  Ohio 171-174 

XXIII.  Supposed  Microsaurian  Species  of  Uncertain  Relationship 175-177 

XXIV.  The  Temnospondylous  Amphibia  from  the  Coal  Measures  of  North  America 178-197 

XXV.  The  Stereospondylous  Amphibia  from  the  Coal  Measures  of  North  America 198-201 

Bibliography  of  the  Fossil  Amphibia,  with  Especial  Reference  to  the  Amphibia  from 

the  Coal  Measures  of  North  America 202-217 

An  Index  to  the  Bibliography  of  Fossil  Amphibia 218-219 

Index 220-222 

vii 


ILLUSTRATIONS. 


PLATES. 

Page. 

1.  Views  along  Mazon  Creek,  Illinois 12 

2.  Drawing  of  type  specimen  of  Micrerpeton  caudatum  Moodie,  from  the  Coal  Measures  of  Mazon  Creek.  .  52 

3.  Specimens  of  Eumicrerpeton  parvum,  Erpetobrachium  mazonensis,  Erierpeton  branchialis,  Mazonerpeton 

longicaudatum,  and  Amphibamus  grandiceps 58 

4.  (1  and  2)  Vertebrae  of  Spondylerpeton  spinatum  Moodie.    (3)  Type  specimen  of  Mazonerpeton  costatuto 

Moodie.    (4)  Type  skeleton  of  Cephalerpeton  ventriarmatum  Moodie.     (5  and  6)  The  halves  of  the 

nodule  containing  a  practically  complete  skeleton  of  Amphibamus  grandiceps  Cope 60 

5.  (1)  A  reconstruction  of  the  Coal  Measures  branchiosaurian,  Eumicrerpeton  parvum   Moodie,  a  small 

primitive  salamander 64 

(2)  A  restoration  of  the  branchiosaurian,  Mazonerpeton,  based  on  two  specimens 64 

6.  Dendrerpeton  acadianum  Owen.     Mandibles,  parts  of  anterior  extremities,  humerus,  etc 68 

7.  Hylerpeton  dawsoni  Owen.    Mandible,  teeth,  rib,  and  bones  of  anterior  extremity.     Bones  of  pelvis  and 

posterior  limb  and  bony  scales 72 

8.  Fritschia  curtidentata  Dawson.    Bones  of  skull  and  anterior  extremity,  bony  rods  of  belly,  of  pelvis,  and 

posterior  extremity 76 

9.  Hylonomus  lyelli  Dawson.     (1)  maxillae  and  skull  bones;  (10)  sternal  bones;  (2)  mandible;  (3)  humerus, 

ribs,  and  vertebrae;  (4)  posterior  limb;  (5)  pelvis;  (6)  caudal  vertebrae 78 

10.  Hylonomus  latidens  Dawson.    Skull,  portion  of  skeleton,  foot,  scapular,  and  sternal  bones,  humerus  and 

rib,  believed  to  belong  to  this  species.    Erect  tree,  Coal  formation,  of  Nova  Scotia 80 

11.  Hylerpeton  longidentatum  Dawson.    Mandible  and  other  bones.    Erect  tree,  Coal  formation 82 

12.  Smilerpeton  aciedentatum  Dawson.     Mandible,  portions  of  skull,  scales,  and  various  bones.    Erect  tree, 

Coal  formation 82 

13.  Dendrerpeton  oweni  Dawson.  Skull,  mandible,  and  bones  of  anterior  limbs,  posterior  limb,  pelvis,  and 

bony  scales 100 

14.  (1  and  2)  Amphibamus  grandiceps  Cope,  from  the  Mazon  Creek  shales 106 

(3)  Sauropleura  (Colosteus)  scutellata  Newberry,  from  the  Linton  Coal  Measures,  the  first  known  of  the 

Ohio  Coal  Measures  Amphibia;  at  first  ascribed  by  Newberry  to  the  fishes,  but  later  correctly  identi- 
fied by  Cope 1 06 

(4)  Type  of  Diceratosaurus  (Ccraterpeton)  punctolineatus  Cope,  from  the  Linton  Coal  Measures 106 

15.  (1)  Dorsum  of  skull  of  Diceratosaurus  punctolineatus  (Cope),  from  the  Coal  Measures  of  Linton,  ( )hio.  .  1 14 

(2)  Ventral  surface  of  skull  of  Diceratosaurus  punctolineatus  (Cope),  from  the  Coal  Measures  of  Linton  .  .  114 

(3)  Pectoral  girdle  of  Diceratosaurus  punctolineatus  (Cope),  from  the  Coal  Measures  of   Linton 114 

(4)  Cervical  or  anterior  dorsal  vertebra  of  Diceratosaurus  punctolineatus  (Cope),  from  the  Linton  Coal 
Measures 114 

16.  (1)  Type  specimen  of  Diceratosaurus  punetolineatus    Cope II.fi 

(2)  Skull  of  Sauropleura  longidentata  Moodie,  from  the  Coal  Measures  of  Linton,  Ohio 116 

(3)  Mandible  of  Sauropleura  longidentata  Moodie,  from  the  Coal  Measures  of  Linton,  Ohio 116 

(4)  Type  specimen  of  Sauropleura  enchodus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio 116 

(5)  Additional  specimen  of  Diceratosaurus  punctolineatus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio.  1 16 

17.  Type  of  Saurerpeton  lathithorax  Cope 126 

18.  (1)  Type  of  Erpetosaurus  sculptilis  Moodie,  from  the  Cannelton  Shales  of  Pennsylvania 132 

(2)  Skeletal  elements  of  Eryops  sp.  indet.,  from  the  Pittsburgh  Red  Shale  at  Pitcairn 132 

(3)  Amphibian  footprints,  Dromopus  aduncus  Branson,  from  the  Mississippian  shales  of  Giles  County, 
Virginia 1 32 

(4)  Type  of  Thinopus  antiquus  Marsh,  amphibian  footprint  from  the  Devonian  of  Pennsylvania 132 

19.  Type  of  Ctenerpeton  alveolatum  Cope,  from  the  Coal  Measures  of  Ohio 134 

20.  (1)  Skull  of  Erpetosaurus  minutus  Moodie,  from  the  Cannelton  slates  of  Pennsylvania 134 

(2)  Skull  and  anterior  part  of  body  of  Ptyonius  pectinatus  Cope,  from  the  Coal  Measures  of  Linton 1  ^4 

(3)  Skeleton  of  Eosauravus  copei  Williston,  from  the  Coal  Measures  of  Linton.    "The  oldest  known  reptile 
from  North  America"  and  closely  related  structurally  to  the  Microsauria 134 

(4)  Part  of  ventral  scutellation  and  ribs  of  Sauropleura  digitata  Cope,  from  the  Coal  Measures  of  Linton .  134 

viii 


»3- 

24. 

25- 


ILLUSTRATIONS.  IX 

Mandible  of  Macrerpeton  deani  MocKlie,  from  the  Linton  Coal  Measures 136 

Portion  of  the  skull  of  Macrerpeton  deani  Moodie,  possibly  of  the  same  individual  as  the  mandible. 

From  the  Linton  Coal  Measures 1  ie, 

Type  of  Cereariomorphus  parvisquaniis  Cope,  from  the  Linton  Coal  Measures 136 

An  additional  specimen  of  Cereariomorphus  parvisquaniis  Cope,  from  the  Linton  Coal  Measures.      .  136 

Skull  of  Sauropleura  scutellata  Newberry.    From  the  Coal  Measures  of  Ohio 136 

Tooth  of  Mastodonsaurus  sp.  indet.  of  the  Carboniferous  of  Kansas 136 

Tooth  of  Mastodonsaurus  giganteus  Jaeger,  from  the  Triassic  of  Germany.    Introduced  for  compar- 
ison with  the  tooth  from  the  Kansas  Carboniferous 136 

Type  of  Leptophractus  lineolatus  Cope,  from  the  Coal  Measures  of  Linton.    Portions  of  maxilla  and 

mandible  of  left  side  with  teeth 160 

Type  of  Proterpeton  gurleyi  Moodie,  from  the  Coal  Measures  of  Illinois,  near  Danville.    Cervical  of 

an  otherwise  unknown  amphibian 160 

Amphibian  phalanx  from  the  Coal  Measures  near  Breeze,  Illinois,  of  an  unknown  species 160 

Large  rib  of  a  stcrcospondylous  stegocephalan,  otherwise  unknown 160 

Type  of  Cope's  species  Tuditanus  mordax  referred  by  him  to  the  cranium,  on  account  of  the  sculp- 
turing of  the  elements,  now  known  to  be  portions  of  the  interclavicle  and  clavicles  of  Diceratosaurus 

punctolineatus 160 

Skull  of  Baphetes  planiceps  Owen,  from  the  Coal  Measures  of  Nova  Scotia 160 

Ventral  seutella;  of  Ctenerpeton  alveolatum  Cope,  from  the  Coal  Measures  of  Ohio 166 

Left  leg  and  pelvis  of  Ichthycanthus  platypus  Cope,  from  the  Coal  Measures  of  Ohio 166 

Type  specimen  of  Pelion  lyelli  Wyman,  from  the  Coal  Measures  of  Ohio.    Supposed  to  represent  the 

ancestral  form  of  the  Salientia • 172 

Scales  of  Cereariomorphus  parvisquamis  Cope,  a  microsaur  from  the  Ohio  Coal  Measures 172 

Type  specimen  of  Cereariomorphus  parvisquamis  Cope 172 

Photograph  of  type  specimen  of  Erpetosaurus  (Tuditanus)  radiatus  Cope,  from  the  Coal  Measures  of 

Linton 180 

Photograph  of  type  specimen  of  Erpetosaurus  tabulatus  Cope,  from  the  Coal  Measures  of  Linton. . .  180 

Photograph  of  the  impression  of  Stegops  divaricata  Cope,  from  the  Coal  Measures  of  Linton 180 

Type  and  only  known  specimen  of  Micrerpeton  caudatum  Moodie,  a  branchiosaur  from  the  Coal 

Measures  shales  of  Mazon  Creek 180 

Type  specimen  of  Erpetosaurus  tuberculatus  Moodie,  from  the  Ohio  Coal  Measures 182 

Type  of  Macrerpeton  huxleyi  Cope,  from  the  Coal  Measures  of  Ohio 182 

TEXT-FIGURES. 

1 .  Map  of  the  Coal  Measures  in  North  America 9 

2.  Distribution  of  Coal  Measures  Amphibia  in  North  America  11 

3.  Topographical  Map  of  Mazon  Creek  Region 13 

4.  Topographical  Map  of  Linton,  Ohio,  Region 16 

5.  Fossil  Tree  Trunk  in  Position 21 

6.  Generalized  Amphibian  Skull 23 

7.  Alimentary  Canal  of  a  Carboniferous  Salamander 26 

8.  Vertebra  and  Ribs  of  Coal  Measures  Amphibia 28 

9.  Ventral  Scutellae  of  Micrerpeton 3° 

10.  Horny  Armor  of  Hylonomus 3' 

n.  The  Skulls  of  two  Microsaurians :   A.  Eoserpeton  tenuicorne;  B,  Ceraterpeton  galvani 33 

12.  Devonian  Footprint 37 

1 3.  Restoration  of  Micrerpeton 53 

14.  Mazon  Creek  Amphibia:   A,  Eumicrerpeton  parvum;  B,  Amphibamus  thoracatus.  59 

140.  Skeleton  of  Mazonerpeton  longicaudatum °2 

146.  Skeleton  of  Mazonerpeton  costatum 64 

15.  A.  Impression  of  Erierpeton  branchialis °5 

B.  Eumicrerpeton  parvum 65 

C.  Larger  Specimen  of  Eumicrerpeton  parvum (,5 

D.  Skeleton  of  Erpetobraehiutn  mazonensis &3 

E.  Rib  of  Mazonerpeton  costatum °5 

15a.  Type  Material  of  Sparodus 6° 

16.  Obverse  of  Cocytinus  gyrinoides ™ 

16a.  Nearly  Complete  Specimen  of  Cocytinus  gyrinoides <*> 

17.  Pelion  lyelli,  supposed  ancestral  Salicntian "4 

1 8.  Skeletal  Elements  of  Smilcrpeton  aciedentatum ■* 


26. 


X  ILLUSTRATIONS. 

19.  Skull  and  Skeleton  of  Tuditanus  punctulatus 87 

20.  Skull  and  Skeleton  of  Tuditanus  longipes 90 

2  1 .  Skeleton  of  Tuditanus  walcotti :  A,  Body ;  B,  Leg 94 

22.  A.  Outline  of  Skull  and  Cranial  Elements  of  Erpetosaurus  minutus  Moodie,  from  the  Cannelton  Slates  of 

Pennsylvania 99 

B.  Outline  of  Skull  and  Cranial  Elements  of  Erpetosaurus  radiatus  Cope,  from  the  Coal  Measures  of  Linton  99 

C.  Palate  of  Erpetosaurus  (tabulatus?),  from  the  Coal  Measures  of  Linton,  Ohio 99 

D.  Outline  of  Skull  and  Cranial  Elements  of  Erpetosaurus  acutirostris  Moodie,  from  the  Coal  Measures  of 
Linton,  Ohio 99 

E.  Outline  of  Larger  Part  of  Skeleton  of  Odonterpeton  triangularis  Moodie,  from  the  Coal  Measures  of 
Linton,  Ohio 99 

F.  Right  Mandible  of  Erpetosaurus  tabulatus  Cope,  from  the  Linton,  Ohio,  Coal  Measures 99 

G.  Skull  Elements  and  Lateral-line  Canals  of  Erpetosaurus  tabulatus  Cope,  from  the  Coal  Measures  of  Linton  99 

23.  Skull  of  Stegops  divaricata 113 

24.  Microsaurian  Skulls  from  Linton,  Ohio:   A,  Diceratosaurus  lsevis;  B,  Diceratosaurus  robustus 119 

25.  Restoration  of  Eoserpeton 1 24 

26.  Restoration  of  Amphibamus 128 

27.  Skeleton  of  Amphibamus  grandiceps 129 

28.  Probable  Appearance  of  Amphibamus 130 

29.  Skeleton  of  Cephalerpeton 133 

30.  Restoration  of  Ptyonius 140 

31 .  Restoration  of  ffistocephalus 144 

32.  Vertebrae  of  Molgophis  brevicostatus 148 

33.  Fore-limb  of  a  Member  of  the  Molgophidas,  Possibly  Pleuroptyx 152 

34.  A.  Interclavicle  of  Sauropleura  pauciradiata 159 

B.  Left  Clavicle  of  Sauropleura  pauciradiata 1 59 

35.  Skull  and  Skeleton  of  Saurerpelon  latithorax 164 

36.  Mandible  of  Leptophractus  dentatus 169 

37.  So-called  Interclavicle  of  Eury thorax  subkevis 170 

38.  Skeletal  Elements  of  Amblyodon 1 77 

39.  Vertebrae  of  Spondylerpeton  spinatum 1 79 

40.  Mandible  of  Macrerpeton  deani 184 

41.  Vertebras  of  Eosaurus  acadianus:  A,  Oblique  Lateral  View;  B,  Oblique  View;  C,  Posterior  View;  D,  Trans- 

verse Section;  E  and  F,  Microscopic  Sections 188 

42.  Skull  and  Mandible  of  Eobaphetes  kansensis:    A,  Outer  View  of  Mandible;  B,  Portion  of  Skull;  C,  Inner 

Surface  of  Mandible 191 

43.  Footprints  of  Dromopus  agilis 200 


THE  COAL  MEASURES  AMPHIBIA 
OF  NORTH  AMERICA 


CHAPTER  I. 

THE  PROBLEM  OF  THE  AMPHIBIA  FROM  THE  COAL  MEASURES. 

The  Amphibia  from  the  Coal  Measures  of  North  America  present  the  problem 
of  the  origin  of  the  land  vertebrates,  since  the  air-breathing  vertebrates  in  the  Coal 
Measures  of  this  continent  are  the  earliest  known  in  the  western  hemisphere.  The 
difference  in  age  between  the  chief  amphibian-bearing  deposits  of  North  America 
and  Europe  is  not  great,  although  it  has  been  asserted  that  Pholidogaster  and  its 
allied  fauna,  described  by  Huxley  from  Scotland  (331),  is  much  older,  probably 
Mississippian.  It  is  interesting  to  note  that  these  earliest  representatives  of  the 
Amphibia  in  Scotland  are  all  temnospondyles,  of  which  there  are  very  few  represen- 
tatives in  the  Coal  Measures  of  North  America. 

The  forms  so  far  described  from  the  North  American  Coal  Measures  present  a 
very  high  degree  of  development  and  differentiation,  the  earliest  known  species 
being  already  specialized  and  well  adapted  for  various  modes  of  life.  As  far  back 
in  geological  time  as  the  middle  Coal  Measures,  when  the  first  well-defined  forms 
are  known,  environmental  conditions  had  effected  a  wide  diversity  of  structure 
within  the  group.  Thus,  early  in  the  geological  history  of  the  land  vertebrates,  we 
have,  among  the  Coal  Measures  Amphibia,  various  forms  which  had  specialized  into 
strictly  aquatic,  terrestrial,  subterrestrial,  and  arboreal,  or  at  least  partly  arboreal. 
Specialization  had  extended  to  the  loss  of  limbs,  ribs,  and  ventral  armature  in  a 
few  species,  and  to  the  acquirement  of  claws,  running  legs,  or  a  long  propelling 
tail  with  expanded  neural  and  haemal  arches  in  others.  The  forms  range  in  size 
from  small  creatures  less  than  an  inch  in  length  to  large  species  which  must  have 
attained  a  length  of  several  feet.  A  rather  interesting  parallel,  though  of  no  phy- 
logenetic  significance,  can  be  drawn  between  the  Amphibia  of  the  North  American 
Coal  Measures  and  the  reptiles  of  to-day.  The  snakes  are  represented  by  the  limb- 
less, snake-like  forms,  such  as  Ptyonius  and  Phlegethontia.  The  lizards  find  their 
counterpart  in  the  Hylonomidae  and  the  Tuditanida?.  No  known  characters  of 
these  animals  tend  to  ally  them  directly  with  any  known  group  of  fishes,  except 
in  the  most  general  way.  These  facts  all  indicate  a  long  antecedent  history  for  the 
amphibian  group  or  else  a  preceding  period  of  greatly  accelerated  development  of 
which  we  now  know  nothing. 

The  Amphibia  whose  remains  have  been  brought  to  light  from  the  Coal  Measures 
have  hitherto  been  regarded  as  pertaining  to  a  single  order,  the  Stegocephalia, 
characterized  by  the'  completely  roofed-over  cranium  and  a  large  parasphenoid. 
The  writer  (469)  had  previously  assigned  5  suborders  to  the  group :  the  Branchio- 
sauria,  Microsauria,  Aistopoda,  Temnospondylia,  and  Stereospondylia.  All  of 
these  groups  are  represented  in  the  Coal  Measures  of  North  America.  It  has  seemed 
inadvisable,  in  the  light  of  our  present  knowledge  of  the  Amphibia,  to  retain  these 
5  groups  as  suborders,  and,  in  the  revised  scheme  of  classification  which  has  been 

3 


4  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

published  elsewhere  (469),  they  are  given  the  rank  of  orders — all  excepting  the 
Aistopoda,  which  are  now  regarded  by  the  writer  as  specialized  Microsauria. 

The  recent  Caudata  are  possibly  represented  in  the  North  American  Coal  Meas- 
ures by  forms  which  may  be  assigned  tentatively  to  the  Proteida.  Such  forms  as 
Cocytinus  gyrinoides,  Hyphasma  loevis,  and  Erierpeton  branchialis  possibly  represent 
this  group  in  the  Pennsylvanian.  This  relationship  is  based  chiefly  on  the  structure 
of  the  hyobranchial  apparatus  and  on  the  general  structure  of  the  species.  The 
three  above-mentioned  species  are,  however,  very  insufficiently  known,  and  the 
relationship  can  hardly  be  regarded  as  more  than  suggested  by  the  characters  which 
are  at  hand. 

The  Salientia,  or  frogs,  may  possibly  have  their  ancestral  type  in  Pelion  lyelli, 
the  first  known  species  from  the  Linton,  Ohio,  Coal  Measures.  Oddly  enough, 
among  the  hundreds  of  specimens  collected  later  from  this  horizon,  not  a  fragment 
can  be  identified  with  this  species.  The  type  specimen  is  unique,  and  although 
incomplete  its  characters  are  suggestive. 

The  Branchiosauria  are  represented  in  North  America  by  four  species:  Mic- 
rerpeton  caudatum,  Eumicrerpeton  parvum,  Mazonerpeton  longicaudatiim,  and  M. 
costatum.  Three  other  genera  which  occur  in  North  America  have  been  placed 
(642)  in  this  group,  but  they  do  not  belong  there,  for  reasons  given  below.  The 
branchiosaurs  were  salamander-like  in  appearance.  They  were  naked,  with  the 
exception  of  small  ovoid  scales  on  the  back  and  the  chevron-shaped  armature  of 
the  ventral  surface,  the  latter  being  almost  universally  present  among  the  Paleozoic 
Amphibia.  They  were  adapted  for  life  in  the  water  for  at  least  the  early  part  of 
their  existence,  as  is  shown  by  the  possession  of  gills  on  many  of  the  late  Carboniferous 
and  early  Permian  forms  of  Europe.  The  group  is,  without  doubt,  ancestral  to 
the  modern  Caudata.  No  branchiosaurians  have  been  described  elsewhere  from 
so  low  in  the  geological  series  as  those  here  given  and  they  are  the  first  and  only 
evidence  of  the  occurrence  of  the  group  in  the  western  hemisphere. 

The  Microsauria  are  represented  in  the  Coal  Measures  by  numerous  forms  which 
are  usually  characterized  as  lizard-like  animals  with  a  well-developed  ventral  scu- 
tellation.  Other  characters,  such  as  the  possession  of  lateral -line  grooves  on  the 
cranium,  the  arrangement  of  the  cranial  elements,  and  the  condition  of  the  ribs, 
will  be  discussed  further  on.  The  pectoral  arch  is  well  developed  and  is  composed 
of  five  dermal  bones  plus  the  regular  skeletal  elements.  The  skeletal  membrane 
bones  are  sculptured  after  the  manner  of  those  of  the  cranium.  The  bodies  of  the 
animals  were,  in  a  few  cases,  covered  with  scales;  but  most  of  them  appear  to  have 
been  completely  naked,  even  the  ventral  armature  being  absent  in  some  cases. 
The  ventral  scutellation  was  especially  strong  and  highly  developed  in  some  of  the 
forms;  e.g.,  in  the  genera  Saarerpeton  and  Sauropleura.  The  vertebrae  are  uniformly 
of  the  hour-glass  or  notochordal  type.  This  is  so  generally  the  case  that  the 
characters  of  the  vertebrae  and  ribs  are  taken  as  the  chief  diagnostic  characters  of 
the  major  groups.  Various  peculiarities  are  seen  among  the  Microsauria,  such  as 
the  development  of  horns  in  various  genera  which  are,  apparently,  related.  The 
order  seems  to  have  gone  completely  out  of  existence  during  the  early  Permian, 


PROBLEM   OF   THE   AMPHIBIA   FROM   THE   COAL   MEASURES.  5 

and  if  their  descendants  continued  on  as  reptiles,  as  has  been  suggested  (469),  we 
do  not  know  the  intermediate  stages. 

The  Aistopoda  are  without  doubt  specialized  microsaurs,  and,  in  the  opinion 
of  the  writer,  are  not  entitled  to  separate  rank.  Some  of  these  forms  reached  a 
high  degree  of  specialization.  One  American  species  has  the  skeleton  reduced  to  a 
long,  slender  head  and  a  slender  series  of  elongate  vertebrae,  all  other  parts  of  the 
skeleton,  even  the  ventral  armature,  being  absent.  The  proportions  attained  by 
this  species,  Phlegetliontia  linearis  Cope,  recall  those  of  the  coach-whip  snake, 
Zamenis  flagellum  Shaw,  of  the  western  plains.  Some  of  the  so-called  Aistopoda 
have  been  credited  by  Fritsch  with  the  possession  of  peculiar  clasping  organs, 
"Kammplatten."  Newberry  has  written  of  the  discovery  of  similar  structures  in 
the  Ohio  Coal  Measures  (498),  but  the  statement  of  the  actual  association  of  these 
"Kammplatten ' '  needs  confirmation.  Dr.  R.  H.  Traquair  wrote  to  the  author  under 
date  of  April  28,  1909: 

"I  maintain  that  the  association  of  a  bundle  of  'Kammplatten'  with  a  specimen  of  Ophi- 
derpeton  in  the  Bohemian  gas  coal  was  entirely  accidental.  Of  such  pitfalls  the  paleontolo- 
gist has  to  beware  or  serious  mistakes  may  be  the  consequence,  as  has  happened  more  than 
once.  I  must,  however,  publish  a  short  paper  on  the  Kammplatten,  for  I  think  I  know 
what  they  are  now." 

Fritsch,  however,  has  very  clearly  figured  a  nearly  complete  specimen  of  Ophi- 
derpeton  (251,  Bd.  iv)  as  possessing  the  Kammplatten  in  place  near  the  cloaca, 
where  he  suggests  they  may  have  served  the  function  of  accessory  copulatory 
organs  or  claspers. 

The  Temnospondylia  are  represented  by  scanty  remains  of  species  from  Illinois, 
Pennsylvania,  and  Nova  Scotia.  The  forms  belonging  to  this  group  are  all  rela- 
tively large,  and  they  had  a  wide  geographical  distribution  during  the  Permian. 
This  group  contains  two  types  of  vertebras,  known  as  the  embolomerous  and  the 
rachitomous,  both  of  which  are  present  in  the  Coal  Measures.  Such  forms  as 
Eosa urns,  Baphetes,  Eobaphetes kansensis,  Macrerpeton,  and  Dendrerpeton  are  regarded 
tentatively  as  temnospondyles,  but  there  is  no  definite  assurance  that  they  are  such. 
It  is  possible  that  Eosaurus  is  a  stereospondyle,  but  the  species  is  too  incompletely 
known  for  a  definite  statement  to  be  made.  The  close  resemblance  between  the 
vertebras  of  Eosaurus  and  Anthracosaurus  has  been  noted  by  Huxley  (332). 

The  Stereospondylia  are  very  scantily  represented  in  the  Coal  Measures,  if  at 
all.  Eosaurus  may  belong  here  as  indicated  above.  The  tooth  and  cranial  frag- 
ments discovered  and  described  by  Williston  from  the  Coal  Measures  of  Kansas 
may  represent  a  stereospondyle  as  he  states  (608),  but  the  evidence  is  incomplete. 
A  fragment  of  a  large  rib  (plate  22,  fig.  4)  of  a  species  from  Linton,  Ohio,  otherwise 
unknown,  may  be  a  stereospondyle.  We  would  expect  an  early  development  for 
this  group,  but  it  is  an  interesting  fact  that  no  stereospondyles  are  known  defi- 
nitely before  the  Triassic,  during  which  period  they  had  an  extensive  distribution. 


CHAPTER  II. 

HISTORY  OF  THE  DISCOVERY  OF  AMPHIBIA  IN  THE  COAL  MEASURES. 

Sir  William  Logan,  in  1841,  discovered  in  the  Coal  Measures  of  Horton's  Bluff, 
Nova  Scotia,  some  tracks  of  Amphibia  which  he  carried  to  London  and  which  Sir 
Richard  Owen  pronounced  to  be  undoubted  "reptilian"  tracks.  This  fact  was 
published  in  1842  (380)  and  was  the  first  recorded  evidence  of  the  occurrence  of 
land  vertebrates  in  the  Carboniferous  rocks  of  the  world.  To  these  tracks  Sir 
William  Dawson  later  gave  the  name  of  Hylopus  logani. 

Two  years  later  Dr.  Gergens  (291)  wrote  a  letter  to  Professor  Bronn,  the  founder 
and  one  of  the  editors  of  the  "Neues  Jahrbuch  fur  Mineralogie,  Geologie  und  Pale- 
ontologie,"  in  regard  to  an  important  discovery  in  the  Carboniferous  rocks  of  Ger- 
many. The  letter  is  of  such  exceptional  interest  in  connection  with  the  history  of 
the  fossil  Amphibia  that  it  is  given  here : 

"In  dem  Brandschiefer  von  Munsterappel  in  Rhein-Bayern  habe  ich  in  vorigen  Jahre 
einen  Salamander  aufgefunden  und  Hrn.  H.  v.  Meyer  in  Frankfurt  zur  naheren  Unter- 
suchung  und  Beschreibung  tibergeben; — Gehort  dieser  Schiefer  der  Kohlen-Formation  ? — 
in  diesem  Falle  ware  der  Fund  in  anderen  Hinsicht  interessant." 

The  form  discovered  by  Dr.  Gergens  and  described  by  Hermann  von  Meyer  as 
an  amphibian  is  a  little  puzzling  as  to  its  characters.  Miall  (449,  p.  183)  says  that 
the  remains  are  too  imperfect  for  close  definition.  The  form,  as  figured,  resembles 
an  immature  branchiosaurian,  as  one  is  at  once  reminded,  from  an  examination  of 
Von  Ammon's  Branchiosaurus  caducas  (7,  Taf.  iv,  fig.  1).  In  1844  Dr.  Alfred 
King  (356)  announced  the  discovery  of  "reptilian"  footprints  in  the  Carboniferous 
of  Pennsylvania. 

The  next  announcement  of  fossil  Amphibia  was  made  by  Goldfuss  (296),  who 
in  1847  described  the  famous  Archegosaurus  from  the  upper  Carboniferous  of  Ger- 
many, from  the  remains  which  had  as  long  ago  as  1777  been  regarded  as  a  fish.  Two 
years  later  Isaac  Lea  (371)  announced  to  the  British  Association  for  the  Advance- 
ment of  Science,  through  Buckland,  the  discovery  of  footprints  in  the  old  Red 
Sandstone  (Mauch  Chunk)  of  Pennsylvania.  These  objects  occur  not  rarely  in  the 
Mauch  Chunk  shales,  which  are  of  upper  Mississippian  age.  Barrell  (21,  p.  460) 
records  the  finding  of  imperfect  tracks  in  the  same  beds,  and  Rogers  (Geology  of 
Pennsylvania,  pt.  11,  1856,  p.  831)  records  three  unnamed  varieties  from  2,200  feet 
below  the  top  of  the  Mauch  Chunk.  Branson  (50)  has  recorded  the  finding  of 
other  amphibian  footprints  from  the  Mississippian  of  Giles  County,  Virginia. 

Lyell  and  Dawson  (396),  in  1853,  read  a  paper  before  the  Geological  Society  of 
London,  in  which  they  announced  the  discovery  of  remains  of  Amphibia  in  the  Coal 
Measures  of  North  America,  although  Dawson  had  previously,  in  1850,  discovered 
the  skull  of  Baphetes  planiceps  Owen,  which  was  not  described  until  the  latter  part 
of  1853  (509).  The  specimen  had  lain  unnoticed  in  the  collection  of  the  Geological 
Society  for  more  than  two  years.    When,  however,  the  announcement  was  made 

6 


HISTORY   OF   DISCOVERY.  7 

by  Lyell  and  Dawson  of  the  discovery  of  Amphibia  in  the  Coal  Measures  of  Nova 
Scotia,  so  much  interest  was  excited  that  the  skull,  now  known  as  Baphetes  plani- 
ceps,  was  brought  to  light  by  the  president  or  secretary  and  was  described  (509) 
by  Sir  Richard  Owen.  The  only  other  known  evidences  of  land  vertebrates  in  the 
Paleozoic  of  North  America,  up  to  this  time,  had  been  the  footprints  described  by 
Lea  and  King  from  the  Mississippian  (Mauch  Chunk)  and  Pennsylvanian  of  Penn- 
sylvania. The  specimens  presented  to  the  Geological  Society  of  London  by  Lyell 
and  Dawson  were  found  at  the  South  Joggins,  Nova  Scotia,  and  consisted  of  scutes, 
a  few  limb  bones,  a  fragment  of  a  jaw,  and  a  few  vertebrae,  a  part  of  which  were 
associated.  The  remains  were  found  quite  accidentally  and  unexpectedly  by  them  in 
the  petrified  trunks  of  ancient  Sigillariae  which  were  exposed  on  the  coast.  Dr. 
Jeffries  Wyman,  of  Harvard  College,  had  examined  these  remains  in  the  United 
States  and  had  pronounced  (638)  them  to  be  amphibian,  comparing  them  with 
similar  elements  in  Menobranehus.  On  the  arrival  of  the  specimens  in  England  they 
were  submitted  to  Sir  Richard  Owen,  who  suggested  the  name  (514)  Dendrerpeton 
acadianum  and  compared  the  remains  with  Archegosaurus.  At  the  same  meeting 
of  the  London  Geological  Society,  Owen  read  a  paper  on  a  small  amphibian  (508) 
from  the  British  Carboniferous  which  he  named  Parabatrachus.  Subsequent  dis- 
coveries have  shown,  however,  that  this  form  belongs  among  the  fishes.  At 
the  meeting  of  the  Geological  Society  held  in  the  latter  part  of  the  same  year  Owen 
announced  (509)  further  discoveries  in  the  Nova  Scotia  coal  beds. 

Hermann  von  Meyer  (436),  in  1857,  described  numerous  stegocephalian  remains 
from  the  upper  Carboniferous  of  Germany.  Dr.  Jeffries  Wyman,  in  the  same  year, 
described  (639)  a  new  form  of  amphibian  from  Linton,  Ohio.  This  form  he  called 
Raniceps  lyelli,  but  as  the  name  Raniceps  had  been  preoccupied  by  Cuvier  for  a 
genus  of  gadid  fishes,  Wyman  later  (1868)  changed  the  name  to  Pelion.  This  was 
the  first  form  to  be  described  from  the  locality  at  Linton,  which  has  since  yielded 
the  remains  of  half  a  hundred  species. 

Dawson  (204),  in  1859,  made  a  further  contribution  to  the  fauna  of  Nova  Scotia 
by  the  description  of  Hylonomus  and  other  species  of  Dendrerpeton  from  the  South 
Joggins  deposits.  Huxley  (331),  in  1862,  described  the  genera  Loxomma  and  Plioli- 
dogastcr  from  the  Carboniferous  of  Scotland.  The  same  year  Owen  made  a  further 
contribution  (514)  to  the  fauna  of  the  Nova  Scotia  beds,  and  Huxley  (332)  discussed 
the  anatomy  of  Anthracosaurus  from  Scotland.  Marsh  (404),  in  the  next  year, 
described,  as  an  enaliosaurian,  the  interesting  Eosaurus  acadianus  from  the  Nova 
Scotia  Coal  Measures,  basing  the  species  on  two  vertebrae,  apparently  from  the 
dorsal  region.  The  vertebrae  resemble  the  stereospondylous  type,  and  Huxley  (332) 
called  attention  to  the  similarity  of  these  vertebrae  to  those  of  Anthracosaurus. 

Cope  (105),  in  1865,  began  his  researches  among  the  Coal  Measures  Amphibia 
of  North  America  by  the  description  of  Amphibamus  grandiceps  from  the  Mazon 
Creek  shales  of  Illinois.  Ten  years  later  (123)  he  published  a  complete  synopsis  of 
the  Carboniferous  Amphibia  of  North  America,  with  especial  reference  to  the  Linton, 
Ohio,  species,  illustrating  many  of  the  forms  now  known  from  Linton.  Between  the 
years  1865  and  1897,  Cope  published  numerous  papers  (105-177)  on  the  Amphibia 


8  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

of  the  Paleozoic,  and  to  his  researches  is  due  a  large  part  of  our  knowledge  of  these 
forms. 

Great  credit  is  due  Dr.  J.  S.  Newberry  (495,  498)  for  the  enthusiasm  and  interest 
which  his  collections  of  Coal  Measures  Amphibia  exhibit.  He  furnished  Cope  with 
the  majority  of  the  type  material  described  by  him,  and  it  was  through  Dr.  New- 
berry's instrumentality  that  the  "Synopsis  of  the  Extinct  Batrachia  from  the  Coal 
Measures"  (123)  was  published.  The  material  which  Dr.  Newberry  had  collected 
he  took  with  him  from  Ohio  to  Columbia  University,  New  York,  and  a  part  of  his 
collection  still  remains  in  the  geological  collection  of  that  institution,  although  the 
greater  portion  has  been  transferred  to  the  American  Museum  of  Natural  History. 
The  Newberry  collection  forms  the  basis  for  the  larger  part  of  this  memoir. 

Between  the  year  1853  and  the  early  nineties,  Dawson  continued  (200-223) 
his  researches  on  the  Amphibia  of  the  Coal  Measures  of  Nova  Scotia.  His  most 
notable  single  work  (208)  is  "The  Air-Breathers  of  the  Coal  Period,"  published  in 
Montreal  in  1863,  in  which  he  gives  a  complete  account  of  the  forms  then  known 
from  Canada,  attempting  some  restorations.  Since  his  death  there  have  been  no 
new  species  described  from  Canada,  and,  so  far  as  I  can  learn,  there  has  been  no 
further  collecting  at  the  South  Joggins. 

Recently  G.  F.  Matthew  (409)  has  rearranged  the  classification  of  amphibian 
footprints  from  Nova  Scotia.  Jaekel  (347)  has  described  very  fully  the  remains  of 
Diceratosaurus  punctolineatus  (Cope)  from  Linton,  Ohio,  basing  the  new  genus  on  a 
species  described  by  Cope  as  a  member  of  Ceraterpeton.  Hay  (316)  has  added  to 
the  knowledge  of  the  anatomy  of  Amphibamus,  his  most  interesting  contribution 
being  the  detection  of  long,  curved  ribs  in  this  form.  This  character  excludes  the 
species  from  the  order  Branchiosauria  and  shows  the  relationship  of  the  form  to 
the  Hylonomidae  and  the  Microsauria.  Schwarz  (540)  has  described  the  characters 
of  the  vertebrae  and  ribs  of  several  genera  of  the  Coal  Measures  Amphibia  and  has 
(541)  offered  his  views  as  to  the  descent  of  the  Amphibia,  based  entirely  on  his  work 
on  the  vertebras  of  species  from  North  America  and  Europe. 

Since  1908  the  writer  has  published  several  contributions  (457-489)  on  the 
Amphibia  from  the  Coal  Measures  of  North  America.  The  results  of  these  investi- 
gations are  given  in  this  work. 


CHAPTER 


STRATIGRAPHIC  AND  GEOGRAPHIC  DISTRIBUTION   OF  AMPHIBIA  IN   THE  COAL 

MEASURES  OF  NORTH  AMERICA. 

There  are  but  four  localities  in  North  America  which  have  furnished  any  notable 
remains  of  Amphibia  in  the  Coal  Measures.  These  are,  in  the  order  of  their  dis- 
covery, the  deposits  at  the  South  Joggins,  Nova  Scotia;  the  Linton,  Ohio, Coal  Meas- 
ures; the  Mazon  Creek,  Illinois,  shales;  and  the  Cannelton  slates  near  Cannelton, 
Pennsylvania.  There  are,  however,  several  other  localities  on  the  continent  which 
have  furnished  evidences  of  Amphibia  in  the  Coal  Measures.  The  principal  one  of 
the  latter  localities  is  doubtfully  of  Coal  Measures  age,  although  recent  discoveries 
would  tend  to  show  it  is  such.  The  deposits  in  question,  those  of  the  Clepsydrops 
shales  of  Vermilion  County,  Illinois,  have,  heretofore,  been  regarded  as  Permian, 
but  the  discovery  of  similar  remains  in 
rocks  of  undoubted  Pennsylvanian  age  in 
Pennsylvania  would  seem  to  indicate  that 
the  Illinois  deposits  were  contemporaneous 
with  them. 

(a)  The  deposits  in  Vermilion  County, 
Illinois,  lie  along  the  north  bank  of  Salt 
Fork  Creek,  at  the  tip  of  the  "Horseshoe 
Bend,"  about  2  miles  south  of  Oakwood, 
Illinois.  They  were  discovered  by  Dr.  J.  C. 
Winslow,  of  Danville,  in  1 875.  The  remains 
discovered  by  him  were  forwarded  to  Pro- 
fessor Cope  for  identification.  Later  the 
deposits  were  thoroughly  explored  by  W.  F. 
E.  Gurley,  and  the  specimens  collected  by 
him  are  now  preserved  (86)  in  Walker  Mu- 
seum, University  of  Chicago.  In  1907,  the 
writer,  while  working  for  the  University  of 
Chicago,  in  exploring  the  same  locality, 
exhausted  the  beds  so  far  as  they  could  at 
that  time  be  uncovered  from  the  landslide 
which  had  overwhelmed  them.  The  for- 
mation in  which  the  bones  occur  is  a  soft  gray 
or  reddish  shale,  and  it  lies  without  any  ap- 
parent stratigraphic  break  on  shales  of  Penn- 
sylvanian age.  Below  these  shales  are  sev- 
eral feet  of  limestone  containing  invertebrates  of  typical  Pennsylvanian  fades.  There 
are  indications  of  at  least  3  species  of  Amphibia  in  the  deposits.  Case  (86)  has  in- 
dicated with  doubt  a  fourth  species.    The  species  are:    Cricotus  Iwteroclitus  Cope, 

9 


Pig.  i.— Map  of  Upper  Pennsylvania!)  showing  land  and 
water  conditions  under  which  the  Coal  Measures 
amphibian  fauna  lived.  It  will  be  noted  that  the 
chief  deposits  which  have  furnished  amphibian  re- 
mains are  on  the  margins  of  the  heavily  shaded 
areas.     (After  Schuchert.) 

Explanation  of  symbols:  Lands  are  white.  Water  areas 
are  lined.  Formation  outcrops  are  black  or  dotted. 
Known  shore-lines  are  solid  lines;  probable  ones 
broken.  Vertical  lines  in  middle  of  continent  incii- 
cate  Gulf  marine. 


10  THE   COAL   MEASURES   AMPHIBIA    OF    NORTH    AMERICA. 

Cricotus  gibsonii  Cope,  Diplocaulus  salamandroides  Cope.  The  remains  are  very 
fragmentary,  and  consist  for  the  most  part  of  incomplete  vertebrae,  with  a  few  small 
skull  fragments. 

(b)  In  1897  Dr.  Williston  (607)  described  some  fragments  of  Cricotus  from  a 
deposit  in  Cowley  County,  near  Winfield,  Kansas.  There  has  been  some  dispute  as 
to  the  age  of  the  deposit,  but  the  consensus  of  opinion  seems  to  be  that  the  beds  are 
of  approximately  the  same  age  as  those  of  Illinois  and  Pennsylvania  in  which  similar 
remains  are  found,  and  those  deposits  are  looked  upon  as  Upper  Pennsylvanic 
(Case  (94),  pp.  239-240).  No  new  forms  were  described  from  Winfield,  since  only  a 
few  fragments  were  obtained.  Williston  referred  the  phalange,  the  fragment  of  a 
jaw,  and  the  tooth  to  Cricotus  heteroclitus  Cope. 

(c)  Later  in  the  same  year  Williston  (608)  announced  the  discovery  of  a  tooth 
of  typical  labyrinthodont  structure  from  near  Louisville,  Kansas  (plate  21,  fig.  6). 
The  tooth  was  accompanied  by  fragments  of  bone  and  was  probably  not  far  from 
the  bed  in  which  it  was  fossilized.  Williston  states  that  the  remains  were  from  the 
shales  which  are  "nearly  at  the  upper  part  of  the  Carboniferous,  probably  within 
one  hundred  feet  of  the  Manhattan  Limestone." 

(d)  In  1894  Marsh  (406)  and  earlier  (1873)  Mudge  (490)  described  footprints  of 
vertebrates  from  the  stone-quarries  near  Osage  City,  Kansas.  The  stone  in  which 
they  were  found  was  a  fine-grained  limestone  which  occurs  near  the  middle  of  the 
Kansas  Coal  Measures. 

(e)  Two  years  later  Marsh  (407)  announced  the  discovery  of  traces  of  the  oldest 
known  (Devonian)  air-breathing  vertebrate.  The  footprints  of  Thin  opus  aiitiuuuswere 
regarded  by  Marsh  as  "apparently  amphibian."  This  still  remains  the  oldest  geo- 
logical evidence  of  air-breathing  vertebrates,  although  Lohest  some  years  ago  (381) 
called  attention  to  remains  from  the  Devonian  of  France  which  he  thought  might  be 
amphibian.  The  footprint  described  by  Professor  Marsh  was  "found  in  the  town 
of  Pleasant,  one  mile  south  of  the  Allegheny  River,  Warren  County,  Pennsylvania, 
by  Dr.  Charles  E.  Beecher,  who  presented  it  to  Yale  Museum,  and  also  furnished 
the  information  in  regard  to  its  geological  position.  ,  .  .  The  geological  horizon 
is  near  the  top  of  the  Chemung,  in  the  upper  Devonian.  In  the  same  beds  are 
ripple  marks,  mud  cracks,  and  impressions  of  rain  drops,  indicating  shallow  water 
and  shore  deposits." 

(f)  Among  the  collections  of  the  American  Museum  there  is  an  impression  of  a 
small  amphibian  foot  obtained  from  Phoenix  Tunnel,  Pennsylvania.  The  impres- 
sion is  in  hard  black  slate  very  similar  to  the  slate  of  the  Cannelton  region.  It  is 
possible  that  the  specimen  may  have  been  obtained  from  the  Cannelton  beds,  since 
they  would  be  expected  to  occur  at  Phoenix  Tunnel.  The  impression  is  rather  small. 
It  is  the  footprint  of  a  5-toed  animal,  probably  of  the  right  foot,  since  no  amphibian 
(465)  so  far  is  known  from  the  Coal  Measures  with  5  digits  on  the  hand.  The  first 
digit  is  short  and  thick,  with  a  large  ball  at  its  base.  The  foot  measures  from  the 
posterior  edge  of  the  palm  to  the  tip  of  the  longest  digit  12  mm.  The  length  of  the 
first  digit  is  7  mm.  The  impression  differs  in  some  respects  from  the  impressions  so 
far  known  from  the  Coal  Measures,  but  no  attempt  will  be  made  to  assign  it  to  a 


STRATIGRAPHIC   AND   GEOGRAPHIC   DISTRIBUTION. 


I  I 


species.  It  may  have  been  made  by  either  a  branchiosaurian  or  a  microsaurian,  but 
more  probably  the  latter,  since  we  do  not  know  of  any  of  the  former  animals  from 
the  Cannclton  beds,  or  in  fact  from  any  of  the  Coal  Measures  beds  excepting  the 
Mazon  Creek  shales.    The  specimen  is  No.  2872  of  the  American  Museum. 


2. 

3- 

4- 
5- 


Fie;.  2. — Distribution  of  Coal  Measures  Amphibia  in  North  America. 

I.  Linton,  Ohio,  near  Yellow  Creek  P.  O.,  Jefferson  County,  Ohio,  on  the  banks  of  Yellow  Creek,  near  the  Ohio  River,  16 
miles  north  of  Steubenville. 

Mazon  Creek  shales,  Grundy  County,  Illinois,  near  Morris. 

"Clepsydrops  shales,"  Salt  Fork  Creek,  Vermilion  County,  Illinois,  near  Oakwood,  on  Tate  farm,  8  miles  west  of  Danville, 
Illinois. 

Danville,  Illinois,  coal  where  the  type  of  Prolerpelon  gurleyi  Moodie  was  found. 

Breeze,  Illinois,  where  Dr.  J.  A.  Udden,  in  1907,  found  a  fragment  of  an  amphibian  phalange  on  the  dump  of  the  Cooper- 
ative Coal  Company. 

6.  Pitcaim,  Pennsylvania,  15  miles  east  of  Pittsburgh. 

7.  Cannelton,  Pennsylvania,  Beaver  County,  Cannelton  slates,  Kittanning  formation,  45  miles  northwest  of  Pittsburgh. 

8.  Fairfield,  Iowa,  where  Dr.  J.  A.  Udden  found  remains  attributed  by  Dr.  Eastman  to  Pkuroptyx  davatus  Cope. 

9.  Louisville,  Pottawatomie  County,  Kansas,  where  Dr.  S.  W.  Williston  discovered  remains  at  Maslodonsaurus  in  the  Coal 

Measures. 
Washington  County,  Kansas,  source  of  type  of  Eobapheles  kansensis  Moodie,  from  the  Coal  Measures. 
Osage  City,  Osage  County,  Kansas,  amphibian  footprints  from  the  Coal  Measures. 
Winfield,  Kansas,  source  of  Cricotus  material. 
Lander,  Wyoming,  in  Wind  River  Carboniferous. 

Pictou,  Pictou  County,  Nova  Scotia,  84  miles  northeast  of  Halifax.    Source  of  Baphetes  planiceps  Owen. 
Joggins  (Joggins  Mines),  Cumberland  County,  Nova  Scotia,  4  miles  from  River  Hebert.    Source  of  HyUrpcton  and  Den- 
drerpeton  faunas. 
16.  South  Joggins,  Nova  Scotia,  source  of  the  Eosaurus  acadianus  Marsh. 


10. 
11. 
12. 

13- 
14. 

15. 


(g)  Dr.  J.  A.  Udden,  in  1907,  discovered  a  fragment  of  a  phalanx  of  some 
amphibian  (plate  22,  fig.  3)  on  the  dump  of  the  Cooperative  Coal  Company,  a  mile 
east  of  Breeze,  Illinois.  It  was  obtained  from  below  the  Shoal  Creek  limestone 
and  somewhere  above  the  (Illinois)  Coal  No.  6,  according  to  Dr.  Udden's  notes.  The 
maximum  width  of  the  phalanx  is  10  mm.  and  it  probably  had  a  length  of  16  mm. 


12  THE    COAL    MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

(h)  Mr.  N.  H.  Brown,  in  191 4,  discovered  in  the  Carboniferous  rocks  to  the  east 
of  the  Wind  River  Mountains,  near  Lander,  Wyoming,  a  single  fragment  of  an 
amphibian.  The  writer  was  accompanying  Mr.  Brown  at  the  time  of  the  discovery 
and  there  can  be  no  doubt  that  the  fragment  was  amphibian ;  the  location  of  the  beds 
was  such  that  no  later  age  than  the  Coal  Measures  can  be  assigned  to  them. 

(i)  Dr.  J.  A.  Udden  (577),  in  1912,  announced  the  discovery  of  an  amphibian 
in  the  Des  Moines  formation  of  Iowa.  These  remains  were  identified  by  Dr.  Charles 
Eastman  as  Plenroptyx  clavatus  Cope.  Since  the  Des  Moines  is  probably  nearly 
contemporaneous  with  the  Mazon  Creek  shales  of  Illinois,  the  discovery  does  not 
extend  the  geological  range  to  any  extent,  but  is  of  interest  as  it  adds  another  note 
to  our  knowledge  of  the  geographical  distribution  of  the  Amphibia  in  the  Coal 
Measures. 

(j)  The  Gurley  collection  of  the  University  of  Chicago  possesses  a  single  cervical 
vertebra  of  some  amphibian  ( ?) .  The  vertebra  is  unlike  anything  previously  described 
and  represents  a  new  form  (plate  22,  fig.  2)  which  may  be  designated  Proterpeton 
gurleyi,  new  genus  and  species.    The  material  was  collected  near  Danville,  Illinois. 

(k)  Deposits  have  been  discovered  in  Pennsylvania  in  which  are  found  the 
remains  of  amphibians  and  reptiles,  very  similar  to  those  from  Vermilion  County, 
Illinois,  Cowley  County,  Kansas,  and  the  Texas  Permian.  The  remains  (plate  18, 
fig.  2)  were  found  in  a  thin  stratum  below  the  "Ames"  limestone,  and  are  therefore 
in  the  Coal  Measures,  fairly  well  below  the  top.  The  fossils,  as  described  by  Case 
(94) ,  consist  of  fragments  which  he  ascribes  to  pelycosaurian  reptiles  and  to  temno- 
spondylous  amphibians.  The  genus  Eryops  (94)  is  recognized  in  several  fragments 
and  a  nearly  complete  dorsal  vertebral  centrum.  Other  types  of  Amphibia  are  like- 
wise represented. 

(I)  The  ironstone  nodules,  in  which  the  Mazon  Creek  fossils  (plate  1)  occur,  are 
found  in  the  shale  which  forms  the  roof  of  the  Morris  or  "No.  2"  Coal  of  Illinois, 
which  "lies  probably  somewhat  lower  than  the  horizon  of  the  Lower  Kittanning 
Coal  of  Pennsylvania"  (599).  "The  nodules  of  iron  contained  in  the  Coal  shales 
on  the  banks  of  Mazon  Creek  near  Morris,  Illinois,  generally  contain  organic 
nuclei,  and  thousands  of  beautiful  specimens  have  been  obtained  there.  They  are 
usually  fragments  of  fern  fronds,  but  are  sometimes  shells,  crustaceans,  myriapods, 
scorpions,  spiders,  cockroaches,  .  .  .  fishes"  (498,  p.  214),  and  amphibians,  of  which 
10  species  are  at  present  known. 

These  species  have  been  arranged  zodlogically  according  to  the  following  plan : 

Class  Amphibia  Linne',  1758. 

Subclass  Euamphibia  Moodie,  1909. 

Order  Branchiosauria  Lydekker,  1889. 

Family  Branchiosanridce  Fritsch,  1879. 
Micrerpeton  caudatum  Moodie,  1909. 
Eumicrerpeton  panmm  Moodie,  1910. 
Mazonerpeton  longicaudatum  Moodie,  191 2. 
Mazonerpeton  costatum  Moodie,  191 2. 
Order  Caudata  Dum£ril,  1806. 
Suborder  Proteida  Cope. 

Family  Cocytinidw  Moodie,  191 2. 

Erierpeton  branchialis  Moodie,  191 2. 


MOODIE 


VIEWS  ALONG  MAZON  CREEK,  ILLINOIS. 

1.  A  nodule  weathering  out  of  the  shale,  at  the  head  of  the  hammer.  Most  of  the  nodules 
at  the  so-called  "lower  beds"  contain  specimens  of  Xeuropteris. 

I.  The  nodules  in  the  creek  bed  at  the  "upper  beds."  Many  of  them  have  been  cracked 
open  by  the  frost. 

3.  Looking  south   at    the    "upper   beds."     The    nodules   found   in    the    background   are 

non-fossiliferous. 

4.  Nodules    may    be  seen  through  the  clear  water  embedded  in  the  shale.     Neuropterid 

insects  in  the  water. 

5.  Looking  for  nodules  at  the  "upper  beds."     The  uppermost  reaches  of  the  .ossiferous 

beds  correspond  with  the  extreme  background  of  the  picture. 
Nodules  in  the  stream  bed  at  the  "lower  beds."     Many  of  these  a,e  cracked  open  by 
the  frost  and  good  specimens  are  sometimes  found  in  the  noduU>. 


S. 


STRATIGRAPHIC   AND  GEOGRAPHIC  DISTRIBUTION.  1 3 

Class  Amphibia  Linne,  1758— Continued. 
Subclass  Lepospondylia  Zittel,  1887. 
Order  Microsauria  Dawson,  1863. 

Family  Amphibamidce  Cope,  1875. 

Amphibamus  grandiceps  Cope,  1865. 
Amphibamus  thoracatus  Moodie,  191 1. 
Cephalerpeton  ventriarmatum  Moodie,  19 12. 
Family  Molgophidm  Cope,  1875. 

Erpetobrachium  mazonensis  Moodie,  191 2. 
Subclass  Stegocephala  Cope,  1868. 

Order  Temnospondylia  Zittel,  1887. 
Suborder  Embolomeri  Cope,  1885. 
Family  Cricotidm  Cope,  1884. 

Spondylerpeton  spinatum  Moodie,  19 12. 

It  will  be  seen  from  the  above  arrangement  that  nearly  all  of  the  orders  of 
Amphibia  are  represented  in  the  Mazon  Creek  fauna.  These  animals  are  the  oldest 
known  land  vertebrates  of  North  America. 


Contour  interval  SOfeet 


Fig.  3. — Portion  of  the  "Morris  sheet"  of  the  U.  S.  Geological  Survey,  to  show 
topography  and  situations  of  the  exposures  of  fossil-bearing  shales  at  Mazon 
River,    a,  the  "Bartlett  place,"  the  so-called  "upper  beds";  b,  "lower  beds." 

The  writer  was  able,  during  July  191 1,  to  spend  a  week  studying  the  fossil  beds 
(479)  at  Mazon  Creek.  The  object  of  the  visit  was  primarily  to  collect  Amphibia, 
but  although  several  thousand  nodules  were  examined,  not  one  contained  an 
amphibian  nor  a  fragment  of  one.  Mr.  J.  C.  Carr,  of  Morris,  Illinois,  who  has  collected 
at  Mazon  Creek  for  more  than  30  years,  has  never  collected  an  amphibian.  These 
facts  interested  me  in  making  the  following  comparison:  If  we  take  100,000  nodules 
as  a  basis  for  computation  of  the  rarity  of  the  various  forms,  something  like  the 
following  will  be  the  approximate  result  of  the  investigation : 


14  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

Of  100,000  nodules,  20,000  will  be  barren  or  contain  only  indeterminate  fragments ; 
68,500  will  contain  plants;  7,500  will  contain  insects,  Crustacea,  myriapods,  scor- 
pions, spiders,  and  other  arthropods;  3,900  will  contain  fish  coprolites  or  scales;  95 
may  contain  fish  or  fragments  of  fish ;  4  may  contain  mollusks ;  and  1  may  contain 
an  amphibian  or  a  fragment  of  one. 

Perhaps  even  100,000  is  low  as  a  basis  of  estimate.  Mr.  Carr  was  of  the  opinion 
that  1  nodule  in  every  500,000  might  contain  an  amphibian. 

The  beds  from  which  the  nodules  are  usually  collected  occur  along  both  banks 
and  in  the  bottom  of  the  creek,  in  two  localities.  One  locality  known  as  the  Bartlett 
place  is  situated  8  miles  southeast  of  Morris,  in  Grundy  County,  Illinois,  Wauponsee 
Township,  N.W.  quarter,  section  30,  Township  33,  Range  8,  the  land  being  now 
owned  by  Mrs.  Emma  Akerly,  of  Wilmington,  Illinois. 

The  fossil-bearing  nodules  occur  throughout  6  to  8  feet  of  shale  along  both  banks 
of  the  creek  at  the  "upper  beds"  (plate  1 ,  fig.  3) ,  as  the  Bartlett  place  is  called.  They 
may  also  be  seen  in  the  bed  of  the  creek,  when  the  water  is  low  (plate  1,  fig.  4), 
still  embedded  in  the  shale.  With  a  potato  fork  the  shale  is  easily  turned  and  the 
nodules  come  out  like  potatoes.  One  sometimes  finds  a  "pocket"  of  nodules  from 
which  as  many  as  a  peck  may  be  secured.  Nearly  every  nodule  has  a  fossil  at  the 
"upper  beds,"  but  all  of  the  fossils  are  not  well  preserved,  possibly  only  1  or  2  out 
of  every  10  being  worth  carrying  to  the  museum.  The  nodules  crack  best  when  wet, 
and  it  requires  some  skill  to  crack  them  evenly.  They  seem  quite  light  and,  in 
one  place  where  the  stream  curves,  are  piled  in  a  long  windrow.  On  this  were  found, 
in  nodules  cracked  by  the  frost,  several  good  crustaceans  and  many  good  plants. 

Table  of  Pennsylvanic  Formations. 

gerjc  Northern  Appalachian.  Bituminous, 

Pennsylvania-Ohio.  Illinois. 

Monongahela  or  Upper  Productive  Coal  Measures {Break""  Fork'i- 

Conemaugh  or  Lower  Barren  Ames  limestone  near  middle: 

(*)   (Pitcairn) Coal  No.  6. 

.  — .  I  Kreeport_ 

Allegheny  or  Lower  Coal  Measures 


t 

c 

3 

w 

o 

« 

c 

£ 

> 

>>< 

i 

o 

c 

0 

c 

(U 

H< 

i 

w    1) 

£3 

£*| 

(*)  (Cannellon)  Coal  No.  2  (*). 
Kittanning  (*).         (Morris?)  (Mazon  Creek). 

(Linton). 
Clarion. 

Homewood. 

Mercer. 

Conoquenessing. 

Sharon. 

(*)  marks  the  position  of  the  Amphibian-producing  horizons  in  these  regions.     (After  Schuchert.) 

The  fossils  at  the  "upper  beds"  are  localized  into  special  strata.  At  one  place 
in  the  upper  part  of  the  deposit,  in  a  reddish  shale,  one  finds  that  insects  are  more 
abundant  than  they  are  lower  down.  The  Crustacea  seem  to  come  from  appar- 
ently the  same  shale.  At  the  lower  end  of  the  deposit  certain  definite  species  of 
Pecopteris  are  localized.  It  is  an  interesting  fact  that  one  seldom  finds  a  Neurop- 
teris  at  the  "upper  beds."  The  most  abundant  fossils  are  the  various  species  of 
Pecopteris  and  Aiimilariu.  When  specimens  of  Neuropteris  are  found  they  are 
usually  discovered  at  the  lower  end  of  the  exposures.  In  one  place  behind  the 
"island"  very  blue  nodules,  hard  and  flinty  and  with  sometimes  well-preserved 


STRATIGRAPHIC   AND   GEOGRAPHIC   DISTRIBUTION.  1 5 

specimens  of  Pecopteris,  are  found  quite  definitely  localized.  These  nodules  are  apt  to 
assume  an  irregular  shape.  These  localizations  of  the  fossils  are,  of  course,  what  we 
would  expect  from  our  knowledge  of  the  recent  fauna  and  flora.  There  is,  to  be  sure, 
more  or  less  intermingling  of  the  species.  The  myriapods,  so  far  as  they  have  been 
found,  are  also  localized.  Mr.  Carr  found  3  within  a  space  of  a  few  feet,  but  again 
these  are  found  widely  scattered.  The  exposures  at  the  "upper  beds"  are  about 
a  quarter  of  a  mile  long.    They  disappear  under  a  heavy  ledge  of  sandy  limestone. 

At  the  ' '  lower  beds' '  (plate  1 ,  fig.  6) ,  those  further  down  the  creek,  conditions  are 
quite  different  from  those  just  described,  although  of  the  same  horizon;  the  banks 
of  the  creek  are  higher  and  almost  perpendicular,  so  that  the  chances  of  collection 
from  the  shales  are  fewer.  The  bed  of  the  creek,  however,  is  wider  and  there  are 
more  nodules  washed  out.  The  most  abundant  fossil  at  this  place  is  Neuropteris. 
The  nodules  at  the  upper  end  of  the  exposure  are  all,  almost  without  exception, 
barren  of  fossils.  The  exposures  here  are  of  about  the  same  extent  as  the  "upper 
beds,"  though  the  species  are  not  so  varied.  Judging  from  the  collections  made 
while  there,  Arthropoda  are  more  abundant  at  the  "lower  beds." 

Bradley  (Geol.  Surv.  Illinois,  iv,  196,  1870)  mentions  the  occurrence  of  these 
nodules  at  or  near  Morris.  Other  than  these  places  the  nodules  have  been  thrown 
out  of  a  coal  mine  near  Braidwood,  Illinois.  Doubtless  close  search  would  reveal 
other  localities  where  the  shale  is  cut  through  in  mining.  The  beds  at  both  places 
are  slightly  folded.  This  is  true  especially  of  the  ' '  upper  beds,"  where  a  conspicuous 
fold  caused  the  beds  to  disappear  in  the  bed  of  the  creek  and  to  reappear  farther 
down  stream.    This  is  directly  across  the  large  "ox-bow"  bend  of  the  creek. 

The  beds  at  Mazon  Creek  were  first  explored  in  1857  by  Mr.  Joseph  Evans, 
who  sent  his  specimens  to  Berlin,  Germany,  where  they  excited  great  interest.  It 
was  he  who  collected  the  type  specimen  of  Amphibatnus  grandiceps  Cope.  Since 
the  time  of  Mr.  Evans  many  have  collected  at  Mazon  Creek,  and  without  doubt 
the  fossil-bearing  nodules  from  this  locality  are  more  widely  scattered  in  the 
museums  of  the  world  than  are  organic  remains  from  any  other  one  horizon. 

So  far  as  we  know  there  was  no  upland  vertebrate  life  at  that  time.  The  forms 
at  present  known  were  confined  to  the  water  or  the  margins  of  the  water.  The 
absence  of  knowledge  of  upland  and  terrestrial  deposits  of  this  time  doubtless 
accounts  for  the  absence  of  known  vertebrates.  It  is,  however,  especially  interesting 
to  speculate  on  the  ancestral  types  of  the  land  vertebrates,  and  it  must  be  admitted 
that  the  Coal  Measures  Amphibia  as  at  present  known  throw  the  ancestry  of  land- 
living  vertebrates  far  back  into  geological  time. 

(w)  The  Cannelton  slates  of  Beaver  County,  Pennsylvania,  have  furnished  3 
species  of  Amphibia  and  fragments  of  other  species  are  represented  in  the  U.  S. 
National  Museum  (462).  The  species  so  far  known  are:  Tuditanus  minimus 
Moodie,  Erpetosaurus  sculptilis  Moodie,  Erpetosaurus  minutus  Moodie. 

They  are  the  first  evidence  of  the  occurrence  of  amphibian  remains  in  these 
deposits.  The  Cannelton  specimens  are  found  in  a  thin  stratum  of  slate  which  forms 
part  of  the  Middle  Cannelton  Coal.  The  Cannelton  slate,  in  which  the  fossils  occur, 
forms  the  roof  of  the  Middle  Kittanning  Coal,  which  is  only  20  to  30  feet  above  the 


16 


THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 


Lower  Kittanning  bed  (I.C.  White),  so  it  becomes  evident  that  the  deposition  of  the 
Cannelton  slates  was  at  only  a  slightly  later  period  than  that  of  the  shales  in  which 
the  Mazon  Creek  nodules  occur,  since  the  Mazon  Creek  shales  form  the  roof  of  the 
Morris,  which  "is  probably  somewhat  lower  than  the  Lower  Kittanning  of  Pennsyl- 
vania." From  the  Cannelton  slates  are  known  the  remains  of  plants,  insects,  Crus- 
tacea, especially  "Eurypterids  found  in  shale  immediately  below  the  Darlington 
(Upper  Kittanning)  Cannel  Coal,  near  Cannelton,  Darlington  Township,  Beaver 
County,  Pennsylvania,  Horizon,  Allegheny  River  Series"  (Hall,  1884).  In  these 
shales  occur  also  fishes  and  the  3  species  of  amphibians  referred  to  above.  The 
Amphibia  known  from  this  region  are  small,  the  largest  of  them  not  exceeding  6 
inches  in  length. 


Contour  interval  200  fret 


Fig.  4. — Portion  of  "West  Virginia-Ohio-Pennsylvania,  Wellsville  Quadrangle"  of  the 
U.  S.  Geological  Survey,  to  show  topography  and  situation  of  Linton  coal  mines.  Some 
fossil  amphibians  doubtless  came  from  across  the  line  in  Columbiana  County. 

(«)  The  Linton,  Ohio,  beds  outcropped  near  Linton  post-office,  which  was  for- 
merly located  at  the  mouth  of  Yellow  Creek,  a  few  hundred  yards  from  the  present 
station,  Yellow  Creek,  Salem  Township,  Jefferson  County,  in  the  valley  of  Yellow 
Creek,  near  the  Ohio  River,  and  thus  near  the  Pennsylvania  state  line. 

In  regard  to  the  exact  location  of  the  town  of  Linton,  which  has  long  since  been 
abandoned,  I  quote  from  a  letter  from  Dr.  Louis  Hussakof,  who  visited  the  locality : 

' '  The  locality  appears  to  have  been  known  as  Yellow  Creek  for  many  years  past.  That 
is  the  name  used  in  the  Geological  Map  of  Ohio  published  by  Orton  in  1888  and  which  was 
based  on  the  earlier  maps  of  Newberry  (1869  and  1879).  When  I  visited  the  place  in  1905, 
and  asked  for  Linton  (which  I  had  not  been  able  to  locate  on  any  map  then  available  to 
me),  hardly  anyone  knew  of  such  a  locality.  Only  one  old  man  in  Steubenville,  Ohio, 
recalled  that  Yellow  Creek  was  identical  with  Linton. 


STRATIGRAPHIC  AND  GEOGRAPHIC  DISTRIBUTION.  1 7 

"Yellow  Creek  is  not  a  village,  but  only  a  R.  R.  station  (on  the  Pennsylvania  R.  R.), 
and  marks  a  spot  where  once  was  an  active  and  prosperous  mine.  Probably  at  a  former 
day  there  was  a  small  post-office  somewhere  in  the  neighborhood  known  as  Linton.  I  did 
not  take  any  photographs,  as  I  was  not  certain  of  the  spot,  or  the  mine,  from  which  the 
fossils  had  come.  There  are  some  cement  mines  within  a  few  minutes'  walk  of  the  station,  but 
no  coal  appears  to  be  mined  at  present  at  Yellow  Creek.  'Smith's  Pit,'  the  coal  mine  best 
remembered  by  the  younger  men,  is  not  worked. 

"Now  as  to  the  question  whether  some  of  the  Amphibia  might  have  come  from  localities 
in  Columbiana  County.  I  believe  it  very  probable  that  they  did.  I  walked  along  the  road 
from  Yellow  Creek  (Jefferson  County)  to  Wellsville  (Columbiana  County),  a  distance  of 
about  2  or  2.5  miles,  and  the  country  seemed  quite  the  same.  Everywhere  one  sees  outcrops 
of  coal  in  the  cuts  along  the  road.  Furthermore,  I  inclose  a  copy  of  a  page  in  an  old  note- 
book of  Professor  Newberry  from  which  you  will  see  that  Coal  Measure  fossil  localities  were 
known  not  only  at  Yellow  Creek,  but  also  from  near  Wellsville.  There  can  be  hardly  a 
doubt  that  most  of  the  specimens  you  have  are  from  Yellow  Creek;  and  quite  a  number  are 
those  collected  by  Sam  Huston." 

Newberry  says,  in  regard  to  the  fauna  of  the  Linton  Coal: 

"The  Linton  locality  is  especially  interesting  and  instructive.  It  has  already  (1889) 
yielded  more  than  20  species  of  fishes  and  nearly  40  species  of  aquatic  amphibians,  all 
inhabitants  of  the  same  body  of  water.  These  were  found  in  a  thin  stratum  of  cannel  which, 
over  a  limited  area,  underlies  a  thick  bed  of  cubical  coal  (No.  6  of  the  Ohio  reports),  of  which 
the  place  is  near  the  top  of  the  Lower  Coal  Measures.  At  Linton,  ...  we  have  evidence 
that  the  great  marsh  in  which  the  peat  accumulated  that  formed  coal  No.  6  was  for  a  time 
a  lake  or  a  lagoon,  inhabited  by  the  fishes  and  amphibians  to  which  I  have  referred.  .  .  . 
Many  of  the  fishes  and  the  amphibians  were  highly  carnivorous  and  powerful,  as  we  leam 
from  their  teeth  and  coprolites.  The  largest  of  the  amphibians  must  have  been  8  or  10 
feet  in  length,  having  strong  jaws,  set  with  numerous  lancet-shaped  teeth  an  inch  or  more 
in  length.  .  .  .  After  a  sufficient  time  had  elapsed  for  many  generations  of  fishes  and 
aquatic  salamanders  to  live  and  die,  the  lake  was  filled  by  the  extension  of  its  peaty  shores 
into  it — just  as  so  many  lakelets  are  filled  and  obliterated  at  the  present  time — and  after- 
ward over  the  cannel  was  formed  a  mass  of  peat,  which  has  now  become  a  stratum  of 
cubical  coal  7  feet  in  thickness. 

' '  In  the  Linton  cannel  are  buried  fragments  or  entire  individuals  of  all  the  inhabitants 
of  this  body  of  water  which  had  hard  parts,  bones,  scales,  spines,  or  teeth,  capable  of  preser- 
vation. Hence  we  get  a  locally  complete  picture  of  the  life  of  the  Carboniferous  age,  and  we 
find  it  to  be  unexpectedly  rich  and  varied.  In  that  age  fishes  and  amphibians  were  the  highest 
forms  of  animal  life,  and  the  amphibians  were  comparatively  newcomers  on  the  earth's 
surface.  Yet  they  had  multiplied  and  differentiated  until  this  little  pool  contained  millions 
of  them,  varying  in  length  from  6  inches  to  10  feet  and  curiously  diversified  in  their  forms, 
their  scales,  and  spines,  and  in  the  ornamentation  of  their  enamel-covered  heads"  (498). 

"To  the  paleontologist  there  are  few  places  in  the  world  more  interesting  than  the  Dia- 
mond mine,  at  Linton,  since  here  we  get  such  a  view  of  the  life  of  the  Carboniferous  age  as 
is  afforded  almost  nowhere  else,  and  of  the  great  numbers  of  species  found  there,  not  more 
than  three  or  four  have  been  met  with  elsewhere"  (497). 

On  page  18  is  a  list  of  the  Amphibia  which  are  thus  far  described  from  the  Lin- 
ton deposits.  They  all  belong,  so  far  as  known,  to  the  Microsauria,  the  reference  of 
any  of  the  species  to  other  orders  being  doubtful.  The  larger  Amphibia  seem  to  be 
indicated  by  a  large  rib  which  resembles  very  much  that  described  by  Huxley  in 
1863  for  Anthracosaurus. 


1 8  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

Amphibia  from  the  Linton  Beds  (51  Species). 
Brachydectes  newberryi  Cope.     Fragment  of  a  skull. 
Cercariomorphus  parvisquamis  Cope.     Impression  of  body. 
Cocytinus  gyrinoides  Cope.    A  skull  and  anterior  dorsal  vertebra?. 
Ctenerpeton  alveolatum  Cope.     Large  portion  of  skeleton,  no  skull. 
Diceratosaarns  Icevis  Moodie.     Complete  skull. 
Diceratosaurus  punctolineatus  Cope.      Anterior  vertebrae,  part  of  skull,  with  ribs  and  portion 

of  ventral  armature. 
Diceratosaurus  robustus  Moodie.     Incomplete  cranium. 
Eoserpeton  (Ceraterpeton)  tenuicome  Cope.     Incomplete  skull. 
Erpetosaurus  acutirostris  Moodie.     Complete  skull. 
Erpetosaurus  obtusus  Cope.     Incomplete  skull. 
Erpetosaurus  radiatus  Cope.     Incomplete  skull. 
Erpetosaurus  tabulatus  Cope.     Incomplete  skull,  with  clavicles. 
Erpetosaurus  tuberculatus  Moodie.     Incomplete  skull. 

Eurythorax  subkevis  Cope.    A  single  interclavicle.    (Operculum  of  lung  fish,  Sagcnodus.) 
Hyphasma  laevis  Cope.    Incomplete  skull  and  anterior  vertebras. 
Ichthycanthus  ohiensis  Cope.     Portion  of  dorsal  region. 
Ichthycanthus  platypus  Cope.     Posterior  portion  of  body. 
Leptophractus  dentatus  Moodie.    Mandible. 
Leptophractus  lineolatus  Cope.     Incomplete  skull. 
Leptophractus  obsolctus  Cope.     Portions  of  skull. 
Macrerpeton  deani  Moodie.     Mandible  and  part  of  skull. 
Macrerpeton  huxleyi  Cope.     Part  of  cranium. 
Molgophis  brevicostatus  Cope.     Part  of  vertebral  column  with  ribs. 
Molgophis  macrurus  Cope.    Vertebral  column. 
Molgophis  wheatleyi  Cope.     Part  of  skull  with  2  5  vertebrae. 
Odonterpeton  triangularis  Moodie.      Skull  and  anterior  part  of  body. 
GLstocephalus  rectidens  Cope.     Part  of  mandible. 
CEstocephalus  remex  Cope.    Skull  and  anterior  part  of  body. 
Pelion  lyelli  Wyman.     Cranium,  fore  part  of  body,  hind  limb. 
Phlegethontia  linearis  Cope.     Skull  and  anterior  part  of  body. 
Phlegethontia  serpens  Cope.     Series  of  22  dorsal  vertebras. 
Pleuroptyx  clavatus  Cope.     Part  of  vertebral  column  and  limbs. 
Ptyonius  marshii  Cope.     Part  of  skull  and  anterior  vertebrae. 
Ptyonius  nummijer  Cope.     Skull  and  greater  part  of  vertebral  column. 
Ptyonius  pectinatus  Cope.     Many  specimens,  some  nearly  perfect. 
Ptyonius  serrula  Cope.     Nearly  complete  skeleton. 
Ptyonius  vinchellianus  Cope.     Skull  and  anterior  vertebra;. 
Saurerpeton  latithorax  Cope.     Skull  and  fore  part  of  body. 
Sauropleura  digitata  Cope.     Greater  part  of  body  minus  skull. 
Sauropleura  (Anisodexis)  enchodus  Cope.     Part  of  jaw. 
Sauropleura  joveata  Cope.    A  single  interclavicle  with  impression. 
Sauropleura  longidentata  Moodie.     Incomplete  skull  with  mandible. 
Sauropleura  newberryi  Cope.     Two  incomplete  skulls  with  vertebras. 
Sauropleura  pauciradiata  Cope.     Elements  of  a  pectoral  arch. 
Sauropleura  scutellata  Newberry.     Imperfect  skeleton. 
Stegops  divaricata  Cope.     Nearly  complete  skull. 
Thyrsidium  fasciculare  Cope.     Dorsal  vertebrae. 
Tuditanus  brevirostris  Cope.     Skull  and  anterior  vertebrae. 
Tuditanus  longipes  Cope.     Part  of  vertebral  column  with  limbs. 
Tuditanus  punctulatus  Cope.     Skull  and  anterior  part  of  body. 
Tuditanus  walcotti  Moodie.     Skull  and  portions  of  body. 

Besides  the  above-listed  species  there  are  others  indicated  by  fragments  too 
poorly  preserved  to  be  worthy  of  specific  designation.  The  Linton  Amphibia  are  all 
apparently  confined  exclusively  to  that  locality.  Species  from  the  Cannelton  slates 
have  been  assigned,  however,  to  genera  which  occur  at  Linton,  i.e.,  Erpetosaurus  and 
Tuditanus.  This  reference  may  be  due  to  lack  of  knowledge,  as  the  forms  are  insuf- 
ficiently known.    A  single  Linton  species  has  been  assigned  to  Ichthyerpeton,  a  genus 


STRATIGRAPHIC   AND  GEOGRAPHIC   DISTRIBUTION.  19 

known  otherwise  only  from  the  Coal  Measures  of  Kilkenny,  Ireland.  Cope  referred 
species  from  Linton  to  the  genus  Ceraterpeton  of  Huxley,  from  Kilkenny,  Ireland,  but 
Jaekel  (347)  and  the  writer  (462)  have  shown  that  the  species  were  incorrectly  assigned 
to  the  genus  Ceraterpeton,  and  that  in  fact  they  represent  widely  distinct  genera.  A 
single  species  has  been  identified  by  Eastman  from  the  Des  Moines  limestone  of  Iowa 
as  identical  with  one  from  Linton,  Pleuroptyx  clavatus  Cope.  The  Linton  fauna  is 
distinct  from  that  of  the  Mazon  Creek  beds,  and  also  from  that  of  South  Joggins, 
Nova  Scotia. 

(0)  The  deposits  in  Nova  Scotia  have  been  correlated  with  the  Coal  Measures 
strata  of  the  United  States  (Bell,  Summ.  Rpt.  Geol.  Surv.  Canada,  1912,  1914,  360- 
371).  They  are  very  near  the  same  age  as  the  Linton  beds  and  come  in  near  the  base 
of  the  Allegheny  River  series.  The  exposures  are  at  the  South  Joggins,  along  the  sea- 
coast.  Here  in  strata  of  clay  interstratified  with  coal  are  found  the  erect  stumps  of 
the  Sigillariae,  and  it  was  in  the  rock  within  these  stumps  that  Lyell  and  Dawson,  in 
1853,  discovered  the  remains  of  the  amphibians  which  they  termed  "reptiles." 

"The  bones  of  Dcndrcrpeton  hitherto  found,  as  well  as  those  of  the  smaller  species,  have 
been  obtained  from  the  interior  of  erect  Sigillariae,  and  all  of  those  in  one  of  the  many  beds 
which,  at  the  Joggins,  contain  such  remains.  The  thick  cellular  inner  bark  of  the  Sigillaria 
was  very  perishable ;  the  slender  woody  axis  was  somewhat  more  durable ;  but  near  the  sur- 
face of  the  stem,  there  was  a  layer  of  elongated  cells,  or  bast  tissue  of  considerable  dura- 
bility, and  the  outer  bark  was  exceedingly  dense  and  indestructible.  Hence  an  erect  tree, 
partly  imbedded  in  sediment,  and  subjected  to  the  influence  of  the  weather,  became  a  hol- 
low shell  of  bark.  When  they  remained  open  for  a  considerable  time,  they  would  constitute 
pitfalls  into  which  animals  walking  on  the  surface  might  be  precipitated.  When  the  sur- 
face was  inundated  all  such  remains  would  be  covered  and  imbedded  in  the  sediment. 
These  seem  to  have  been  the  precise  conditions  of  the  bed  which  afforded  these  remains." 
(Dawson,  223,  1894.) 

Fifteen  species  have  been  described  from  the  Joggins  deposits.  Two  are  known 
from  the  Albion  mines,  south  Nova  Scotia,  where  were  obtained  the  remains  of 
Baphetes  planiceps  Owen  and  B.  minor  Dawson. 

The  following  17  species  of  Amphibia  are  known  from  the  Carboniferous  of 
Canada : 

Amblyodon  problematicum  Dawson.     Teeth  and  fragments. 
Baphetes  minor  Dawson.     An  incomplete  mandible. 
Baphetes  planiceps  Owen.     An  incomplete  cranium  from  Albion. 
Dendrcrpeton  acadianum  Owen.     A  jaw,  limb  bones,  and  fragments. 
Dendrerpeton  oweni  Dawson.     Phalangeal  bone  and  fragments. 
Eosaurus  acadianus  Marsh.     Two  dorsal  vertebrae. 
Fritschia  curtidentata  Dawson.     A  mandible,  vertebras,  ribs. 
Hylerpeton  dawsoni  Owen.     Mandible,  teeth  and  incomplete  maxilla. 
Hylcrpeton  intermedium  Dawson.    Mandible  and  portions  of  skull. 
Hylerpeton  longidentatum  Dawson.     Fragments  of  mandible  and  skull. 
Hylonomus  lattdens  Dawson.     Mandible  and  teeth. 
Hylonomus  lyelli  Dawson.     Incomplete  skeleton  and  part  of  skull. 
Hylonomus  multidens  Dawson.     Fragments  of  skull. 
Hylonomus  wymani  Dawson.     Mandible  and  vertebrae. 
Platystegos  loricatum  Dawson.     Incomplete  skull,  vertebrae. 
Smilerpeton  aciedentatum  Dawson.     Teeth,  ribs,  fragments. 
Sparodus  sp  indet.     Teeth,  scales. 


20  THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 

(p)  All  the  remains  representing  the  above  species  were  collected  by  Sir  J.  Wil- 
liam Dawson  at  the  South  Joggins  and  at  the  mines  of  Albion,  with  the  exception 
of  Eosaurus,  which  was  collected  by  O.  C.  Marsh.  The  collections  of  Dawson  are 
now  in  the  Peter  Redpath  Museum  of  McGill  University  in  Montreal  and  in  the 
British  Museum  of  Natural  History  at  South  Kensington,  London.  The  history  of 
the  discovery  of  the  deposits  and  their  amphibian  fossils  at  the  South  Joggins  is  so 
interesting  that  it  was  thought  worth  while  to  reproduce  in  large  part  Dawson's 
paper  "On  the  Mode  of  Occurrence  of  Remains  of  Land  Animals  in  Erect  Trees  at 
the  South  Joggins,  Nova  Scotia,"  published  in  1891  in  the  Transactions  of  the  Royal 
Society  of  Canada,  section  iv,  p.  127: 

"The  remarkable  section  of  coal-formation  rocks  at  the  South  Joggins,  in  Cumberland 
County,  has  long  been  known  as  one  of  the  most  instructive  in  the  world;  exhibiting  as  it 
does  a  thickness  of  5,000  feet  of  strata  of  coal-formation  in  a  cliff  of  considerable  height, 
kept  clean  by  the  tides  and  waves,  and  in  the  reefs  extending  from  this  to  the  shore,  which 
at  low  tide  expose  the  beds  very  perfectly.  It  was  first  described  in  detail  by  the  late  Sir 
W.  E.  Logan  (Report  Geol.  Surv.  Canada,  1844),  and  afterwards  the  middle  portion  of  it 
was  still  more  detailed  by  the  author  (Dawson),  more  especially  in  connection  with  the 
fossil  remains  characteristic  of  the  several  beds  and  the  vegetable  constituents  and  accom- 
paniments of  the  numerous  seams  of  coal  (Jour.  Geol.  Soc.  Lond.,  x,  p.  1,  1853).  It  was  on 
occasion  of  a  visit  of  the  author  in  company  with  Sir  Charles  Lyell,  and  in  the  pursuit  of 
these  investigations,  that  one  of  the  most  remarkable  features  of  the  section  was  disclosed 
in  185 1.  This  is  the  occurrence,  in  the  trunks  of  certain  trees  imbedded  in  an  erect  position 
in  the  sandstones  of  Coal-mine  Point,  of  remains  of  small  reptiles,  which  with  one  exception, 
a  specimen  from  the  Pictou  coal-fields,  were  the  first  ever  discovered  in  the  Carboniferous 
rocks  of  the  American  continent,  and  are  still  (1801)  the  most  perfect  examples  known  of 
a  most  interesting  family  of  coal-formation  animals,  intermediate  in  some  respects  between 
reptiles  proper  and  batrachians,  and  known  as  Microsauria.  With  these  were  found  the 
first-known  Carboniferous  land  snails  and  millipedes.  Very  complete  collections  of  these 
remains  have  been  placed  by  the  author  with  his  other  specimens  in  the  Peter  Redpath 
Museum  and  in  the  British  Museum. 

"A  forest  or  grove  of  the  large  ribbed  trees  known  as  Sigillariw  was  either  submerged 
by  subsidence  or,  growing  on  low  ground,  was  invaded  with  the  muddy  waters  of  an  inun- 
dation, or  successive  inundations,  so  that  the  trunks  were  buried  to  the  depth  of  several 
feet.  The  projecting  tops  having  been  removed  by  subaerial  decay,  the  buried  stumps 
became  hollow,  while  their  hard  outer  bark  remained  intact.  They  thus  became  hollow 
cylinders  in  a  vertical  position  and  open  at  the  top.  The  surface  having  then  become  dry 
land,  covered  with  vegetation,  was  haunted  by  small  quadrupeds  and  other  land  animals, 
which  from  time  to  time  fell  into  the  open  holes,  in  some  cases  nine  feet  deep,  and  could 
not  extricate  themselves.  On  their  death,  and  the  decomposition  of  their  soft  parts,  their 
bones  and  other  hard  portions  remained  in  the  bottom  of  the  tree  intermixed  with  any  vege- 
table debris  or  soil  washed  in  by  rain,  and  which  formed  thin  layers  separating  successive 
animal  deposits  from  each  other.  Finally,  the  area  was  again  submerged  or  overflowed  by 
water,  bearing  sand  and  mud.  The  hollow  trees  were  filled  to  the  top  and  their  animal  con- 
tents thus  sealed  up.  At  length  the  material  filling  the  trees  was  by  pressure  and  the  access 
of  cementing  matter  hardened  to  stone,  not  infrequently  harder  than  that  of  the  contained 
beds,  and  the  whole  being  tilted  to  an  angle  of  200,  and  elevated  into  land  exposed  to  the 
action  of  the  tide  and  waves,  these  singular  coffins  present  themselves  as  stony  cylinders 
projecting  from  the  cliff  or  reef,  and  can  be  extracted  and  their  contents  studied.  The  sin- 
gular combination  of  accidents  above  detailed  was,  of  course,  of  very  rare  occurrence,  and. 


STRATIGRAPHIC  AND  GEOGRAPHIC  DISTRIBUTION. 


21 


in  point  of  fact,  we  know  only  one  set  of  beds  at  the  South  Joggins  in  which  such  remains 
so  preserved  occur;  nor  is  there,  so  far  as  I  am  aware,  any  other  known  instance  elsewhere. 
Even  in  the  beds  in  question,  only  a  portion  of  the  trees,  about  15  out  of  30,  have  afforded 
animal  remains.  We  have,  however,  thus  been  enabled  to  obtain  specimens  of  a  number  of 
species  which  would  probably  otherwise  have  been  unknown,  being  less  likely  than  others  to 
be  preserved  in  properly  aqueous  deposits.  Such  discoveries  on  the  one  hand  impress  us  with 
the  imperfection  of  the  geological  record;  on  the  other,  they  show  us  the  singular  provisions 
which  have  been  made  in  the  course  of  geological  time  for  preserving  the  relics  of  the 
ancient  world,  and  which  await  the  industry  and  skill  of  collectors  to  disclose  their  hidden 
treasures. 

"There  is  evidence  in  coprolitic  matter  on  one  of  the  surfaces  within  the  trunks,  and 
also  in  certain  trails  on  these  surfaces,  that  some  of  the  imprisoned  animals  lived  for  a  time 
in  their  subterranean  prisons;  that  they  crept  around  their  walls  in  search  of  a  way  of 
escape,  and  that  the  larger  animals  fed  on  smaller  species  entrapped  along  with  them." 

After  the  discovery  of  these  en- 
tombed amphibians  Sir  William  Dawson 
was  given  a  grant  of  £50  from  the  Gov- 
ernment Fund  by  the  council  of  the  Royal 
Society  of  London,  to  aid  in  the  extrac- 
tion of  these  trees  and  the  collection  of 
their  contents.  The  trees  were  carefully 
taken  out  and  their  contents  examined ; 
the  portions  containing  the  animal  re- 
mains were  carefully  boxed  to  be  taken 
to  Montreal  for  final  cleaning  and  study. 
Erosion  goes  on  rapidly  at  the  South 
Joggins,  but  no  one  has  paid  any  atten- 
tion to  the  occurrence  of  Amphibia  along 
the  coast  of  Nova  Scotia  within  recent 
years. 

(q)  A  deposit  which  will  be  of  un- 
doubted interest  in  connection  with  the 
occurrence  of  Amphibia  in  the  Coal 
Measures  is  that  which  outcrops  along 
the  banks  of  Rock  Creek  in  the  south- 
western part  of  Douglas  County,  Kansas, 


Fig.  5.— Dawson's  tree  No.  13  at  the  South  Joggins,  Nova  Scotia. 
Upper  part,  in  situ,  in  the  reef  after  it  had  been  exposed  by 
blasting.    (After  Dawson,  based  on  a  photograph.) 


in  Marion  Township  (Township  14  south,  Range  18  east,  SW.  and  SE.  quarters  of 
section  7),  about  2  miles  from  the  now-abandoned  post-office  of  Twin  Mounds,  so 
called  from  the  two  flat-topped,  elongated  mounds  of  Oread  limestone  to  the  west 
of  the  town. 

The  interest  in  these  beds  is  not  due  to  the  discovery  of  Amphibia  in  them,  but 
the  possibilities  of  such  discoveries.  This  is  indicated  by  the  occurrence  of  fossils, 
in  nodules  similar  to  those  obtained  from  Mazon  Creek,  which  are  identical  gener- 
ically,  and  in  most  cases  specifically,  with  the  Mazon  Creek  animals  and  plants. 


22  THE   COAL   MEASURES    AMPHIBIA   OF   NORTH   AMERICA. 

The  fossils  so  far  collected  from  this  interesting  locality  are : 

Insecta  (Identified  by  Dr.  E.  H.  Sellards). 

Spiloblattina  maledicta  (Scudder).    The  basal  half  of  a  front  wing. 
Etoblattina  sp.    The  hind  wing  of  a  cockroach. 

Arachnida. 

Anthrocomartus.     Impression  of  the  body. 

Prestwichia  dance  M.  &  W.     Nearly  complete  specimen. 

Crustacea. 

Acanthotelson  stimpsoni  M.  &  W.    Three  nearly  complete  individuals. 

Plants  (Identified  by  Mr.  J.  C.  Carr,  of  Morris,  III). 
Pecopteris  sp. 

Sagittaria  reticulata  Lesquereux. 
Annularia  longijolia  Lesquereux. 
Annularia  inflata  Lesquereux. 
Pecopteris  villosa  Brongniart. 
Neuropteris  decipiens  Lesquereux. 
Pecopteris  serpulifolia  Lesquereux.     By  far  the  most  abundant  plant  is  Pecopteris. 

The  fossils  occur  in  definite  strata  of  nodules  immediately  above  a  io-inch  coal 
seam  which  is  worked  for  local  consumption.  The  coal  lies  near  the  base  of  the 
exposure  in  the  more  western  portion  of  the  outcrop,  but  it  is  raised  by  an  anticlinal 
fold  to  near  the  top  of  the  creek-banks  by  the  bridge  across  Rock  Creek,  along  the 
banks  of  which  the  shales  are  exposed.  Nodules  containing  fossils  are  found  most 
abundantly  at  the  western  exposure  on  the  McKinzie  place,  only  a  few  having  been 
found  near  the  bridge. 

Below  the  coal-seam,  nodules  of  various  shapes  and  sizes  occur,  but  they  seldom 
contain  fossils  and  never  good  ones.  Occasionally,  as  at  Mazon  Creek,  fragments 
of  plants  adhere  to  the  outside  of  the  incrusting  shale.  A  single  nodule  may  have 
adhering  to  it  fragments  of  4  genera  of  plants.  The  fossiliferous  nodules  all  occur 
above  the  coal  and  are  most  prolific  and  abundant  immediately  above  the  seam, 
within  the  first  10  inches.  In  the  same  stratum  of  shale  with  the  nodules  are  found 
abundant  impressions  of  plants  in  the  shale,  often  perfect  fronds  being  uncovered. 
(See,  in  this  connection,  Twenhofel  and  Dunbar,  1914,  "Nodules  with  Fishes  from 
the  Coal  Measures  of  Kansas,"  Amer.  Journ.  Sci.,  xxxvm,  pp.  157-163.) 

G.  F.  Matthew  (408-413)  has  described  numerous  genera  and  species  of  foot- 
prints, presumably  amphibian,  from  the  Carboniferous  of  Canada.  The  impressions 
indicate  small  creatures  for  the  most  part.  Other  imprints  have  been  described  by 
Logan,  Dawson,  Lyell,  Marsh,  Mudge,  and  Lea.  Since  the  present  work  is  intended 
largely  for  a  morphological  review,  only  passing  notice  can  be  given  to  the  ichnites. 
The  literature  on  the  " Ichnites"  has  been  brought  together  in  Hay's  "Bibliography 
and  Catalogue  of  Fossil  Vertebrata  of  North  America,"  pp.  538-553.  References 
since  the  publication  of  Hay's  catalogue  (317)  will  be  found  in  the  bibliography  at 
the  end  of  this  work.  Footprints  are  of  interest  in  that  they  are  the  only  evidence  we 
have  of  the  occurrence  of  land  vertebrates  in  the  Devonian  and  Mississippian  of 
North  America. 


CHAPTER  IV. 

THE  MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA. 

The  anatomy  of  the  Coal  Measures  Amphibia  presents  many  primitive  types  of 
structure.  Their  organization  represents  a  stage  passed  through  in  the  ontogeny 
of  higher  vertebrates.  The  animals  are  similar  in  a  general  way,  yet  so  diverse  are 
the  modifications  which  they  have  suffered  under  different  environmental  conditions, 
that  close  scrutiny  is  needed  to  discern  the  exact  relationship  of  the  forms.  Our 
knowledge  of  this  relationship  is  based  on  the  structures  preserved,  which  are 
largely  skeletal,  since  little  is  known  of  the  soft  anatomy  (471)  of  the  air-breathing 
vertebrates  of  the  Coal  Measures.    The  pubis  is  ossified  in  the  Paleozoic  Amphibia 


a.  com. 


Fig.  6. — Generalized  figure  of  dorsum  of  an  early  amphibian 
skull  to  show  position  of  elements  and  terminology 
adopted  m  this  work.  The  outline  is  based  on  that 
of  Eryops,  but  is  in  no  way  intended  to  indicate  that 
form. 

a.  com,  anterior  commissure  of  lateral-line  canals;  com, 
commissural  communication  between  infra-  and  supra- 
orbital lateral-line  canals;  fr,  frontal;  inf,  interfrontal; 
inn,  internasal;  info,  infraorbital  lateral-line  canal;  it, 
intertemporal;  jl,  jugal  lateral-line  canal;  j,  jugal; 
lar,  lacrimal;  mx,  maxilla;  n,  nasal;  oc,  occiput;  occ, 
occipital  cross-commissure  of  the  lateral-line  system; 
or,  orbit;  par,  parietal;  pof,  postfrontal;  pmx,  premax- 
illa;  pf,  prefrontal;  po,  postorbital;  pp,  postparietal ; 
q,  quadrate;  qj,  quadratojugal ;  spo,  supraorbital 
lateral-line  canal;  sq,  squamosal;  spt,  supratemjxjral ; 
/,  temporal  lateral-line  canal;  tab,  tabulare. 


-  Jl 


later  than  the  ischium  and  ilium;  the  carpus  and  tarsus  are  cartilaginous;  the  verte- 
brae consist  of  a  pleurocentrum  and  two  neurocentra,  thus  paralleling  conditions  in 
modern  mammalian  embryos. 

(a)  The  skull  of  the  Coal  Measures  Amphibia  has  (fig.  6)  essentially  the  same 
structure  in  the  different  groups.  It  is  largely  formed  of  bones  of  intramembranous 
origin,  representing  the  face  bones  of  the  mammalian  skull.  The  skull  in  life  was 
doubtless  a  chondrocranium  with  the  membrane  bones  laid  down  upon  the  carti- 
laginous box  containing  the  sense-organs,  as  in  the  sturgeon  (Acipenser),  where  the 
surface  bones  of  the  face  were  probably  originally  scales,  which  later  became  consol- 
idated into  large  bony  scutes.  The  membrane  bones  of  the  early  Amphibia  may 
have  been  originally  derived  from  scales,  but  at  present  nothing  is  known  of  this 
origin ;  doubtless  the  elements  had  an  intramembranous  origin  in  the  ancestors  of 

23 


24  THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 

the  group.  Judging  from  Credner's  studies  on  the  series  of  specimens  of  Branchio- 
saurus  amblystomus  Credner  (187),  the  skull  bones  do  not  ossify  completely  until 
relatively  late  in  the  life  of  the  individual.  The  skull  in  the  youngest  individual 
figured  by  Credner  {op.  cit.,  Taf.  xvi,  fig.  1)  seems  to  be  largely  cartilaginous,  with 
beginnings  of  separation  into  the  skeletal  elements.  The  manner  and  time  of  devel- 
opment and  ossification  of  the  skull  seems  to  proceed  much  as  it  does  in  modern 
amphibians.  The  condition  found  in  the  skull  of  Cryptobranchus  allegheniensis  or 
Necturus  maculosus  will  represent  pretty  accurately  the  condition  of  most  of  the 
Coal  Measures  Amphibia.  The  face  bones  in  certain  forms  were  sculptured  and 
cut  by  lateral-line  canals. 

A  median  suture  divides  the  skull  into  two  equal  regions  dorsally.  On  the  sides 
of  this  median  suture  lie  pairs  of  elements  which  are  common  to  all  higher  verte- 
brates. These  elements  are:  the  premaxillae,  nasals,  frontals,  parietals,  and  post- 
parietals.  All  of  these  elements  vary  somewhat  in  shape  and  slightly  in  arrange- 
ment, but  always  occupy  the  same  relative  positions.  To  the  side  of  these  elements 
lie  the  prefrontal,  the  postfrontal,  the  supratemporal,  the  squamosal,  and  tabulare, 
and  occupying  the  margin  of  the  skull  are  the  maxilla,  the  jugal,  the  quadratojugal, 
and  possibly  the  quadrate  in  a  few  forms.  The  parietal  foramen  occurs  usually 
within  the  parietal  bone,  but  its  position  is  subject  to  slight  variations  and  it  may 
occur  on  the  suture  between  the  frontal  and  the  parietal,  or  even  far  posterior  near  the 
postparietal.  The  nostrils  often  lie  well  forward  and  are  included  by  the  premaxillae, 
nasals,  and  prefrontals.  The  orbit  is  usually  well  posterior,  but  it  may  occur  far 
forward.  It  is  bounded  by  the  prefrontal,  the  frontal,  the  postfrontal,  the  post- 
orbital,  and  the  jugal.  Sometimes  the  lacrimal  is  present  and  has  been  clearly 
identified  on  the  anterior  margin  of  the  orbit  in  a  few  cases. 

(b)  Sclerotic  plates  often  occur  within  the  orbits,  and  are  not  confined  to  any  par- 
ticular group,  though  they  are  quite  constant  among  the  Branchiosauria.  They  are 
usually  delicate,  thin,  broad  plates  which  evidently  overlap  and  operate  as  in  mod- 
ern animals.  The  number  varies,  as  many  as  30  occurring  within  the  orbit  of  one 
branchiosaur.  Between  the  margin  of  the  orbit  and  the  sclerotic  plates  there  often 
occur,  in  the  Branchiosauria  (186)  particularly,  small  scale-like  particles  which 
were  doubtless  embedded  in  the  heavy  skin  above  the  orbit  during  life. 

(c)  The  palate  of  the  skull  is  very  incompletely  known,  being  indicated  in  a  very 
few  cases.  These  specimens,  however,  show  that  the  characters  of  the  palate  were 
quite  similar,  if  not  identical,  in  essential  respects  with  the  palate  among  the  Euro- 
pean species  of  the  same  or  slightly  later  time. 

A  large  cultriform  parasphenoid  occupies  the  posterior  portion  of  the  palate,  on 
either  side  of  which  in  some  species  lies  the  posterior  palatine  foramen.  On  the 
sides  of  the  anterior  prolongation  of  the  parasphenoid  lie  the  vomers  (186),  membra- 
nous bones  often  bearing  minute  tubercular  teeth,  apparently  adapted  for  crushing. 
The  vomers  and  the  maxillae,  with  sometimes  the  palatine,  surround  the  anterior 
palatine  foramen,  which  is  almost  always  present;  sometimes,  however,  quite  small. 
The  transverse  or  ectopterygoid  unites  the  pterygoid,  a  broad  plate  of  thin  bone, 
with  the  maxilla  and  jugal. 


MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA.  25 

(d)  The  teeth  of  the  Coal  Measures  Amphibia  (194)  are  remarkably  similar  in  the 
various  forms.  They  are  always  slender,  pleurodont  denticles  arranged  in  a  single 
row  on  the  jaws  or  as  tubercular  eruptions  on  the  palate  bones,  with  a  large  pulp- 
cavity  and  the  enamel  often  striated.  The  food  of  the  creatures  must  have  been 
small  Crustacea,  worms,  insects,  and  succulent  vegetation,  such  as  is  the  food  of 
the  modern  Amphibia. 

(e)  The  occiput  is  formed  of  partially  ossified  (465)  ex-  and  basi-occipitals, 
though  these  elements  are  never  firmly  united  by  ossific  union.  Often  a  pair  of 
condyles  occur,  one  on  either  exoccipital.  The  occiput  was  usually,  however,  car- 
tilaginous and  no  trace  of  its  structure  is  preserved. 

(/)  The  mandible  is  usually  as  long  as  the  skull  and  is  slender.  It  is  composed  of 
6  elements  so  far  as  known  (465) ;  these  are  the  articular,  the  surangular,  the  angular, 
the  coronoid,  the  dentary,  and  the  splenial.  Other  elements  may  be  present,  but 
the  anatomy  of  this  portion  of  the  animals  is  not  very  completely  known.  The  bones 
are  sculptured  and  cut  by  lateral-line  canals  (458)  in  a  few  forms.  Whether  the 
articular  operated  on  an  osseous  or  cartilaginous  quadrate  is  unknown,  though 
certain  specimens  seem  to  indicate  an  osseous  condition  for  that  element.  The 
anterior  symphysis  was  doubtless  ligamentous,  the  halves  always  separating  before 
fossilization.  The  dentary  always  bears  a  single  row  of  pleurodont  teeth,  which 
may  vary  greatly  in  size  and  number. 

(g)  The  hyoid  apparatus  is  well  preserved  in  a  few  forms  (123).  Doubtless  it 
was  present  in  all  of  them,  though  it  has  seldom  been  preserved.  The  condition  of 
the  hyobranchial  apparatus  in  Cocytinus  gyrinoides  (text-fig.  16)  from  the  Coal 
Measures  of  Linton,  Ohio,  seems  to  indicate  that  the  species  was  a  perennibranchiate 
salamander  (123).  It  is  well  known  from  the  studies  of  Credner  that  the  Euro- 
pean Branchiosauria,  in  the  young,  possessed  external  branchiae  (187)  supported  by 
lateral  basibranchials.  The  gill-arches  seem  to  have  been  slightly  calcified  or 
ossified  in  a  few  cases,  and  they  supported  denticle-like  projections  which  bore 
the  gill-filaments.  When  the  Branchiosauria  had  attained  a  length  of  100  mm. 
or  more  they  lost  their  gills  (187).  This  change  was  accompanied  by  the  reduc- 
tion of  the  tail,  expansion  of  the  pelvis,  and  increase  in  ossification  of  the  skull 
and  skeletal  elements.  Gills  have  not  yet  been  detected  among  the  American 
Branchiosauria. 

(//)  The  eye  in  a  few  species  had  a  large  amount  of  black  pigment,  as  indicated 
by  the  blackening  of  the  stone  in  the  Mazon  Creek  nodules.  Professor  Cope  (107) 
thought  that  this  would  indicate  a  nocturnal  and  crepuscular  habit  for  these  verte- 
brates, since  the  pigmentum  nigrum  of  the  choroid  is  largely  developed.  Other  than 
this  suggestion  nothing  is  known  of  the  soft  parts  of  the  head. 

(I)  The  alimentary  canal  (text-fig.  7)  is  beautifully  preserved  as  a  cast  in  three 
specimens  of  the  American  branchiosaur  species  Eitmicrcrpcton  parvum  Moodie 
(471)  from  the  Mazon  Creek  beds.  The  nodules  which  contain  these  interesting 
little  fossils  measure  less  than  3  inches  in  long  diameter.  The  fossil  salamanders, 
about  30  mm.  in  length,  are  preserved  on  their  backs  and  occur  as  nearly  as  is 
possible  in  the  center  of  the  nodule. 


26 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


If  it  were  not  for  the  fact  that  the  oesophagus  became  loosened  and  dropped 
from  its  place  shortly  after  death,  the  alimentary  canal  would  be  in  place  and  would 
immediately  recall  a  freshly  dissected  specimen  of  a  recent  salamander.  The  ante- 
rior end  of  the  oesophagus  lies  obliquely  across  the  chest  region  with  its  tip  pointing 
slightly  downward.  The  length  of  the  oesophagus  proper,  in  one  specimen,  is  only 
about  3  mm.  As  it  is  preserved,  the  oesophagus  lies  in  a  semi-sigmoid  curve  with  the 
convexity  anterior,  and  enters  the  cardiac  portion  of  the  stomach  by  a  gradual  con- 
striction. The  stomach  is  clearly  preserved  as  a  distinct  sac-like  organ  with  two 
lobes  which  correspond  to  the  cardiac  and  pyloric  limbs.  It  measures  about  7  mm. 
in  length  by  2  mm.  in  its  greatest  diameter.  The  muscular 
constriction  which  divides  the  organ  into  pyloric  and  cardiac 
divisions  occurs  at  a  distance  of  4  mm.  from  the  upper  end. 
The  pylorus  is  designated  by  a  rather  pronounced  constric- 
tion which  may  be  partly  accidental,  although  it  recalls  the  Is' 
pylorus  of  modern  frogs.  From  this  constriction,  which  lies  >j>, 
on  the  left  side  of  the  fossil,  as  it  is  preserved,  the  duodenal  ^ 
portion  of  the  intestine  makes  a  straight  course  posteriorly 
to  near  the  anal  region,  where  it  takes  a  sharp  bend  and 
curves  back  to  run  parallel  with  itself  for  the  distance  of 
4  mm.  In  its  upward  course  the  intestine  enlarges,  and 
practically  the  same  enlargement  continues  throughout  the 
remainder  of  the  course  to  the  anus.  At  a  distance  of  1 
mm.  from  the  anal  end,  the  rectum  dilates  probably  0.125 
mm.  to  form  the  cloaca.  After  the  intestine  has  continued 
its  parallel  course  for  the  4  mm.,  as  above  stated,  it  turns 
abruptly  to  the  right  for  a  distance  of  2  mm.  It  then  runs 
posteriorly  for  a  short  distance,  then  bends  back  and  under 
itself  to  again  make  a  double  sigmoid  curve,  when  at  a  dis-  pIG. 
tance  of  6  mm.  from  the  anus  it  assumes  a  straight  course, 
which  it  continues  to  the  end. 

The  anus  lies  at  a  level  which  is  approximately  that  of 
the  lower  end  of  the  femur,  which  is  preserved  as  an  impres- 
sion on  the  left  side  of  the  fossil,  thus  agreeing  in  its  position 
with  that  found  in  modern  Caudata.  Lying  inside  the  curve 
of  the  stomach  and  partly  inclosed  by  the  oesophagus  is  a  smooth  area  which  may 
possibly  represent  the  impression  of  some  of  the  accessory  digestive  glands,  such 
as  the  liver.  Occurring  in  this  smooth  area  are  numerous  fine  lines  which  possibly 
represent  the  impressions  of  blood-vessels;  but  they  are  so  imperfectly  preserved 
that  one  can  not  be  sure. 

Representatives  of  several  genera  of  the  modern  Caudata  have  been  dissected 
in  order  to  make  a  direct  comparison  of  the  fossil  alimentary  canal  with  that  of  the 
recent  forms.  The  alimentary  tract  of  Desmognathus  fuscus  Raf.  from  the  vicinity 
of  Ithaca,  New  York,  resembles  in  a  marked  degree  that  of  the  fossil  form.  The 
nearest  approach  to  the  condition  there  represented  is  found,  however,  in  an  imma- 


7. — Alimentary  canal  of  a 
Coal  Measures  salamander  as 
illustrated  by  the  smaller  speci- 
men of  Eumicrerpeton  parvum 
Moodie,  from  the  Mazon 
Creek  shales.  X  3.  Original 
in  Yale  University  Museum. 
a,  anus;  dd,  duodenum;  in, 
intestine;  /,  impression  of 
liver(?);  oes,  oesophagus;  si, 
stomach. 


MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA.  27 

ture  branchiate  individual,  some  47  mm.  in  length,  of  Diemyctylus  torosus  Esch., 
from  a  fresh-water  pond  on  Orcas  Island  in  Puget  Sound.  The  presence  of  this 
species  on  the  island  is  very  suggestive.  It  is  of  extreme  interest,  too,  that  the  con- 
dition represented  in  the  alimentary  tract  of  the  fossil  branchiosaurs  should  resem- 
ble so  closely  that  of  an  immature  rather  than  a  mature  form. 

(j)  The  vertebral  column  is  clearly  and  readily  separable  into  cervical,  dorsal, 
sacral,  and  caudal  regions.  The  neck  is  always  short,  with  from  5  to  10  vertebra?, 
cervical  ribs  often  present.  The  dorsal  region  is  not  long,  but  varies  from  20  to  30 
in  the  constituent  vertebrae.  There  is  a  single  sacral  vertebra  not  always  to  be 
readily  distinguished  from  those  of  the  dorsal  and  anterior  caudal  series.  The  tail 
may  be  very  short  or  extremely  long,  with  neural  and  haemal  spines  elongate  and 
flattened  into  an  oar-like  appendage.  The  distal  caudals  are  in  some  species  car- 
tilaginous, apparently  always  so  in  the  Branchiosauria. 

(k)  The  atlas  and  axis  are  unknown  among  the  American  specimens,  but  we  are 
able  to  infer  from  the  structure  of  the  other  vertebrae  what  this  must  have  been;  and 
our  inferences  are  partly  confirmed  by  the  conditions  existing  in  the  European  forms 
(187).  The  atlas,  apparently,  consisted  of  a  pair  of  neurocentral  plates  which  are 
partly  ossified,  partly  embedded  in  cartilage,  judging  from  the  edges  of  the  plates 
which  have  been  preserved.  The  centrum  seems  not  to  have  been  present  in 
the  atlas,  or  if  present  it  was  only  very  slightly  developed  and  quite  free  from 
the  neural  pieces  and  largely  embedded  in  cartilage.  A  fairly  accurate  picture  of 
the  condition  of  the  atlas  and  axis  may  be  seen  on  examining  a  cow,  pig,  or  chick 
embryo  (378)  in  the  early  stages  of  vertebral  development,  which  has  been  cleared 
by  the  Schultze  method  (Amer.  Journ.  Anat.,  vil,  No.  4,  1908). 

(/)  The  dorsal  vertebrae,  as  well  as  those  of  the  other  series,  present  a  primitive 
character  (fig.  8)  in  the  persistence  of  the  notochord  (540).  Among  the  Branchio- 
sauria the  notochord  was  not  at  all  or  but  slightly  constricted  intravertebrally,  but 
among  the  Microsauria  it  was  constricted  so  far  that  the  notochordal  remnants  in 
each  centrum  resemble  an  hour-glass. 

The  structure  of  the  vertebrae  among  American  forms  agrees  fully  with  that 
outlined  by  Credner,  Fritsch,  and  others  for  the  European  species.  The  details  of 
structure  are  so  fully  given  by  Zittel  (642,  pp.  346~353)  and  by  Schwarz  (540,  541) 
that  it  will  not  be  necessary  to  state  more  here  as  to  their  structure,  since  there  is 
nothing  new  to  add  concerning  the  American  species. 

The  temnospondylous  vertebra  of  the  same  nature  and  type  as  exhibited  by  the 
Permian  forms  has  its  representatives  (94,  478)  in  the  Coal  Measures.  Spondyler- 
peton  spinatum  Moodie  (478)  (plate  4,  figs.  1,  2)  and  Eryops  sp.  (plate  18,  fig.  2) 
indicate  the  embolomerous  and  rachitomous  types  of  vertebral  structure.  The 
occurrence  of  these  widely  different  types  of  vertebral  structure  indicates  a  long  his- 
tory for  the  group  prior  to  the  Coal  Measures.  This  history  is  further  indicated  by 
footprints  in  the  Mississippian  and  Devonian  of  this  continent. 

(m)  The  ribs  (fig.  8)  are  very  diverse  in  structure  and  in  their  mode  of  articula- 
tion (541)  with  the  vertebral  column.  The  characters  of  the  ribs  and  vertebrae  con- 
stitute the  best  means  of  classification  of  these  animals  so  far  discovered.     In  the 


28 


THE   COAL    MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


Branchiosauria  the  ribs  are  always  straight,  heavy,  and  short,  and  articulate  intra- 
vertebrally  upon  a  large  and  strong  transverse  process.  They  occur  throughout 
the  vertebral  column.  There  is  a  single  pair  of  sacral  ribs  which  are  not  to  be  clearly 
distinguished  from  the  pre-sacral  and  post-sacral  series.  The  cervical  and  caudal 
ribs  are  shorter  than  the  dorsal  series.  The  branchiosaurian  rib  is  composed  almost 
entirely  of  perichondral  ossification.  It  presents  the  same  condition  as  does  the 
humerus  of  the  cow  embryo  of  2  to  3  inches  in  length.     The  ribs. of  the  branchio- 


Fig.   8. — Vertebrae  and  ribs  of  Amphibia  from  the  Coal  Measures  of  Linton,  Ohio. 
Originals  in  Geol.  Inst.  Berlin.       (All  after  Schwarz.) 

A.  Caudal  vertebra  of  CRstocephalus  remex  Cope.     Lateral  view.     X  4. 

B.  Caudal  vertebra  of  Ptyonius  vinchellianus  (?)  Cope.    Lateral  view.     X  6. 

C.  Dorsal  vertebra  of  Ptyonius  peclinalus  Cope.    Lateral  view.     X  9. 

D.  Notochordal  cones  and  spinal  canal  of  Thyrsidium  fasciculare  Cope.     X  3. 

E.  Rib  of  Molgophis  sp.  Cope.    X  1.75.     r  =  capitulum;  /  =  tubercuium. 

F.  Vertebra  of  Molgophis  sp.  Cope,  from  ventral  side.    X  2. 

G.  Dorsal  vertebra  of  Ptyonius  pectinatus  Cope.    From  above.     X  8. 
H.  Dorsal  vertebra  of  Thyrsidium  fasciculare  Cope,  from  below.    X  2.5. 

I.  Vertebra  of  Phlegethontia  linearis  Cu\x-,  from  alxive.     X  5.5 

J.  Rib  of  (Estocephalus  remex  Cope,  from  posterior  dorsal  region.    X  5. 

K.  Dorsal  vertebra  of  Thyrsidium  fascicidare  Cope,  from  above.    X  1.5 

L.  Anterior  dorsal  vertebra  (cervical?)  of  Thyrsidium  fasciculare  Cope.    Lateral  view.     X  1.5. 

M.  Vertebra  of  Phlegethontia  linearis  Cope,  from  side.     X  5. 

saurs  are  identical  in  every  way  with  the  ribs  of  modern  salamanders  and  form  one 
of  the  strong  arguments  in  favor  of  the  relationship  of  the  Branchiosauria  to  the 
Caudata.  Among  the  Microsauria  the  ribs  are  always  long,  slender,  curved,  and 
intercentral.  They  may  be  either  single  or  double  headed,  but  usually  the  former. 
They  resemble  in  their  characters  the  ribs  of  some  of  the  early  reptiles  and  an 
attempt  has  been  made  to  relate  the  Microsauria  (469)  to  the  primitive  reptiles  on 
this  basis.  The  ribs  of  the  other  groups  are  still  unknown.  Indeed,  representatives 
of  the  Temnospondylia  and  the  Stereospondylia  are  very  scanty  in  the  American 


MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA.  29 

Coal  Measures.  One  large  rib  (plate  22,  fig.  4)  may  represent  a  labyrinthodont, 
but  nothing  is  known  of  the  species  to  which  the  rib  belonged. 

(h)  The  pectoral  girdle  (187)  is  a  very  simple  and  uniform  structure,  although 
the  details  of  the  association  of  the  elements  still  remain  to  be  determined.  A 
single,  median,  usually  large  and  elongate  interclavicle  occupies  the  ventral  line 
of  the  chest.  This  is  morphologically  the  same  element  which  occurs  in  the  middle 
line  of  the  chest  of  the  lizards.  It  is  a  dermal  bone  and  is  usually,  especially  among 
the  Microsauria  (462),  highly  sculptured.  It  varies  considerably  in  size  and  shape, 
but  is  remarkably  uniform  throughout  the  various  groups.  Lying  anterior  to  the 
interclavicle  and  overlapping  its  antero-lateral  margins  lie  the  two  clavicles,  which 
are  usually  diamond-shaped  and  are  sculptured,  dermal  bones.  The  position  of  the 
coracoid  is  still  uncertain,  and  in  fact  its  clear  association  in  the  pectoral  girdle  of 
these  species  is  still  a  question.  It  seems  to  be  constant  in  the  European  (186,  251) 
species  and  is  usually  represented  by  a  small  rounded  plate  of  bone,  which  in  life  no 
doubt  had  a  large  amount  of  cartilage  to  form  its  borders.  A  cleithrum  (285)  has  been 
ascribed  to  one  of  the  Linton,  Ohio,  species  (plate  15,  fig.  3)  by  Jaekel  (347),  but  this 
needs  confirmation.  An  osseous  scapula  is  usually  present,  resembling  the  scapula  of 
modern  salamanders,  in  that  it  was  largely  embedded  in  cartilage.  The  position 
of  the  pectoral  girdle  is  largely  a  matter  of  doubt,  especially  for  the  American  spe- 
cies. After  death  and  before  fossilization  the  girdle  was  always  moved  by  post- 
mortem shifting,  so  that  its  exact  relation  to  the  ribs  and  vertebral  column  is  still 
in  doubt.  Credner  (186)  has  restored  the  pectoral  girdle  close  behind  the  head, 
which  would  cause  an  amount  of  rigidity  in  the  body  which  probably  did  not  exist. 

(0)  The  arm  consists  of  the  humerus,  radius,  ulna,  and  4  digits.  The  characters 
of  the  arm-bones  are  such  as  is  constant  among  primitive  animals  and  developing 
mammals.  The  osseous  portion  is  perichondral.  Epiphyses  are  totally  lacking  and 
it  is  doubtful  if  the  endochondrium  was  at  all  ossified.  The  digits  are  often  termi- 
nated by  ungual  phalanges,  although  usually  the  terminal  phalanx  was  merely 
embedded  in  the  web  of  the  foot ;  and  among  the  terrestrial  forms  a  claw  was  well 
developed.  An  osseous  carpus  is  not  known  in  the  species  from  the  Coal  Measures. 
Its  impression  indicates  a  broad  hand,  well  adapted  for  swimming. 

(p)  The  pelvic  girdle  consists  uniformly  (462)  of  the  ilium  and  ischium.  A 
small  rounded  pubis  is  present  in  some  of  the  later  forms  of  Amphibia;  it  is,  however, 
totally  absent  from  the  Coal  Measures  species.  The  condition  of  the  pelvis  is 
paralleled  by  the  partially  grown  pelvis  of  mammalian  embryos  in  which  the  elements 
ossify  in  the  order  of  ilium,  ischium,  and  pubis.  The  ilium  is  always  the  larger  of 
the  elements.  It  supported  or  was  attached  to  the  sacral  rib  by  means  of  a  liga- 
mentous union.  The  ischium  did  not  ossify  completely  until  the  animal  was  nearly 
mature.  The  union  between  the  elements  of  the  pelvis  was  probably  of  a  loose, 
membranous  sort  or  else  the  whole  mass  was  embedded  in  cartilage;  of  the  two 
hypotheses  the  former  is  the  more  probable. 

The  pubis  is  indicated  as  a  calcified  quadrangular  plate  in  a  specimen  olAmphi- 
bamus  grandiceps  Cope  (478)  from  the  Mazon  Creek  shales,  and  it  is  present  as  a 
rounded  osseous  element  among  some  of  the  Permian  forms. 


3° 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


(q)  The  leg  (fig.  21,  B)  is  composed  of  the  femur,  tibia,  fibula,  and  5  digits.  The 
tarsus  is  usually  cartilaginous,  a  single  osseous  tarsus  (483,  484)  being  known 
(plate  23,  fig.  1)  from  America.  The  distal  phalanges  may  or  may  not  be  clawed, 
depending  on  the  habits  of  life  of  the  animal.  The  elements  of  the  leg  are  ossified 
in  a  similar  manner  to  those  of  the  arm. 

(r)  The  ventral  scutellation  (fig.  9),  so  commonly  present  among  all  groups  of 
Amphibia  in  the  Coal  Measures,  consists  of  a  series  of  ossifications  or  calcifications 
in  the  myocommata.  Among  modern  amphibians  they  occur  as  thin  perpendicular 
planes  of  connective  tissue  which  are  sometimes  cartilaginous,  especially  in  Necturus, 
regarded  by  Wilder  (Memoirs  of  the  Boston  Society  of 
Natural  History,  vol.  v,  No.  9,  p.  400,  fig.  6,  1903)  and 
by  Wiedersheim  (605,  p.  58)  as  a  homologue  or  predecessor 
of  the  sternum,  although  Wiedersheim  says: 

' '  The  sternum  appears  for  the  first  time  in  Amphibians  in 
the  form  of  a  small  variously  shaped  plate  of  cartilage  situated 
in  the  middle  line  of  the  chest.  It  arises  as  a  paired  cartilaginous 
plate  in  the  inscriptiones  tendineae  of  the  rectus  abdominis 
muscle,  and  therefore  may  be  looked  upon  as  corresponding 
to  a  pair  of  'abdominal  ribs.'  Such  cartilaginous  abdominal 
ribs  must  have  been  present  in  greater  numbers  in  the  ancestors 
of  existing  Urodeles." 

This  supposition  is  fully  sustained  by  the  anatomy  of  FlG 
the  Branchiosauria  (459),  which  must  be  looked  upon  as 
the  actual  ancestors  of  the  Caudata.    Wilder  says  of  these 
structures  in  Necturus  (op.  cit.,  p.  400): 

"The  several  cartilaginous  rudiments  which  represent  this  part  {i.e.,  sternum)  in  Nec- 
turus are  somewhat  difficult  of  detection  and  thus  entirely  escaped  the  attention  of  the 
earlier  investigators.  They  consist  of  a  number  of  thin  cartilages  found  in  several  suc- 
cessive myocommata  of  the  pectoral  region  and  confined  mainly  to  the  area  covered  by  the 
overlapping  epicoracoids. " 

The  homologue  of  the  ventral  scutellce  is  found  in  plesiosaurs,  crocodiles,  Sphenodon, 
and  other  reptiles  in  the  "abdominal  ribs,"  and  the  same  myocommatous  ossifica- 
tions undoubtedly  go  to  the  formation  of  the  chelonian  plastron.  What  the  causes 
were  which  produced  the  development  of  the  ventral  scutellae  to  such  a  high  degree 
among  the  primitive  land  vertebrates  is  uncertain,  but  they  are  certainly  more 
highly  developed  among  the  primitive  reptiles  and  amphibians  than  among  the 
later  members  of  those  classes.  Among  the  Amphibia  of  the  Coal  Measures  they 
attained,  in  some  forms,  a  high  degree  of  development  and  differentiation.  They 
are  present  in  all  families  so  far  known,  except  the  Tuditanidas,  in  which  the  myo- 
commata may  have  been  cartilaginous.  The  Sauropleuridae  present  the  highest 
development  of  these  structures  among  the  American  forms,  in  which  the  scutes  are 
large  and  osseous.  Among  the  Branchiosauria  they  are  calcified  or  partially  ossi- 
fied and  are  always  arranged  en  chevron  on  the  belly,  chest,  arms,  and  throat,  their 
arrangement  and  direction  of  the  chevron  being  modified  according  to  the  myomeres 
of  the  various  regions.  The  ventral  scutellse  of  the  European  Branchiosauria  are 
figured  and  described  fully  by  Credner  (192,  p.  21,  figs.  4  to  11). 


9. — Ventral  scutelke  of  Micrer- 
pclon  caudatum,  a  Coal  Meas- 
ures salamander  from  Mazon 
Creek.  X  5.  /.femur;  /(,  hu- 
merus; Is,  lines  of  scutes;  v, 
vertebral  column. 


MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA.  31 

(s)  Scales  (fig.  10  and  plate  24,  figs.  2  and  3)  are  present  on  the  body  of  (462, 485) 
several  species.  It  is  a  matter  of  regret  that  their  preservation  is  so  imperfect 
that  nothing  can  be  found  out  as  to  their  structure.  The  Linton  species,  which  pos- 
sess scales,  are,  of  course,  carbonized  and  hence  impracticable  for  microscopic  study, 
and  in  the  Mazon  Creek  species  of  Amphibamus  and  Micrerpeton  the  scales  have 
been  replaced  by  kaolin.    The  bodies  of  two  species  (Cercariomorphus  parvisquamis 


a 


Fig.  10. — Horny  armor  of  back  of  Hylonomus.  a,  imbricated  scales;  b,  horny  plates; 
c,  horny  spines  or  tubercles;  d,  small  imbricated  scales.  (After  Dawson,  based  on 
a  photograph.) 

and  Ichthyerpeton  squamosum)  of  the  Linton  Coal  Measures  Amphibia  were  com- 
pletely scaled.  The  scales  in  the  Branchiosauria  (462),  so  far  as  they  have  been 
observed,  are  slightly  imbricated;  rounded,  with  concentric  markings  after  the 
manner  of  the  modern  cyprinoid  fish-scale.  They  are  extremely  minute,  and  whether 
or  not  they  covered  the  entire  body  of  the  animal  is  unknown.  On  the  body  of 
Cercariomorphus  the  scales  have  the  appearance  of  being  tubercular  without 
imbrication,  and  they  apparently  covered  the  entire  bodily  surface  of  the  animal. 


32  THE   COAL  MEASURES  AMPHIBIA  OF   NORTH  AMERICA. 

Among  the  Paleozoic  Amphibia  from  Nova  Scotia  as  described  by  Dawson  and 
Owen  (193,  201,  485)  scales  are  well  developed  and  frequent,  although  the  details 
as  to  their  occurrence  on  the  bodies  of  the  animals  are  still  unknown,  since  the  Nova 
Scotian  species  are  all  based  on  very  fragmentary  remains.  Dawson  (208,  p.  34) 
has  given  a  detailed  discussion  of  the  discovery  and  anatomy  of  the  various  types 
of  scales  possessed  by  the  species  from  the  Coal  Measures  of  Nova  Scotia.  Suffice 
it  to  say  here  that  none  of  the  scales  appear  to  be  bony,  but  have  a  cuticular  appear- 
ance with  concentric  markings.  Some  of  them  are  tubercular,  and  Dawson  thought 
that  a  few  specimens  indicated  that  some  of  the  species  possessed  scaly  lappets  and 
a  dorsal  nuchal  fringe  of  scaly  skin  along  the  back.  He  has  indicated  these  facts 
in  his  restorations  of  the  forms.  The  scales  were  all  carbonized  and  burned  readily 
with  a  strong  flame.  A  section  of  the  scale  shows  a  thick  upper  corium  with  a  vas- 
cular body  (208,  pi.  iv,  fig.  29)  much  like  a  fish-scale.  Fragments  of  the  skin  were 
also  preserved  with  the  scales.    Dawson  says  of  the  skin: 

"One  of  my  specimens  is  a  flattened  portion  of  cuticle  two  and  a  quarter  inches  in 
length.  The  greater  part  of  the  surface  is  smooth  and  shining  to  the  naked  eye,  and  under 
the  microscope  shows  only  a  minute  granulation.  A  limited  portion  of  the  upper  and,  I 
suppose,  anterior  part  is  covered  with  imbricated  scales,  which  must  have  been  membra- 
nous or  horny,  and  generally  have  a  small  spot  or  pore  near  the  outer  margin,  some  having 
in  addition  smaller  scales  or  points  on  their  surfaces"  (208,  pi.  iv,  figs.  22  and  25). 

(t)  Muscle  tissue  (fig.  21)  is  preserved  in  a  single  specimen,  previously  described 
by  the  writer  (464,  p.  17,  pi.  7,  fig.  1).  The  carbonized  muscles  show  a  myomeric 
arrangement  and  the  portions  preserved  probably  represent  one  of  the  recti  muscles 
of  the  abdominal  wall. 

(u)  The  lateral-line  system  in  the  Coal  Measures  Amphibia  will  be  best  under- 
stood from  a  comparative  review  of  the  occurrence  of  these  organs  among  all  extinct 
Amphibia.  Since  all  the  orders  of  Amphibia  are  represented  in  the  Coal  Measures, 
such  a  review  will  not  be  out  of  place  here. 

The  preservation  of  the  lateral-line  system  among  ancient  Amphibia  is  due  to 
the  fact  that  the  skull  of  many  forms  (especially  the  later  and  larger)  are  grooved 
and  marked  by  a  regular  series  of  furrows  and  pits,  in  which  the  sense-organs  of  the 
lateral-line  system  were  contained  (see  fig.  6),  as  well  as  by  the  preservation  of  a 
series  of  clearly  marked  scales  on  the  sides  of  the  tails  and  bodies  of  others.  The 
grooves  are  never  arched  over  as  in  the  Macropetalichthyidae,  where  "in  favorable 
specimens  each  is  shown  to  be  covered  by  a  delicate  roof  perforated  by  two  lines 
of  minute  openings"  (Dean,  N.  Y.  Acad.  Sci.  Mem.,  vol.  11,  pt.  in,  p.  115).  They 
are  always  widely  opened  canals,  either  with  perfectly  smooth  bottoms  and  sides  or 
roughened  with  large  pits,  or  even  becoming  a  series  of  well-marked  pits.  An 
attempt  has  been  made  (458)  to  homologize  the  organs  with  those  of  fishes. 

The  nomenclature  adopted  here  for  the  canals  does  not  depart  from  that  used 
by  Allis  for  Amia  (Journ.  of  Morphology,  vol.  II,  1889).  The  supraorbital  and  infra- 
orbital canals  are  readily  correlated  with  those  of  the  same  name  in  fishes,  where 
they  are  very  clearly  marked.  The  anterior  commissure  is  also  homologous  with 
that  of  the  fishes,  as  is  also  the  canal  here  called  the  "  antorbital  commissure. "    The 


MORPHOLOGY  OF  THE  COAL  MEASURES  AMPHIBIA.  33 

others  arc  not  so  readily  homologized.  The  upper  canal  (see  fig.  6)  in  the  posterior 
part  of  the  cranium  is  here  designated  the  temporal  canal.  It  is,  however,  clearly  a 
part  of  the  infraorbital  of  the  fishes.  Its  relations  in  the  Stegocephala  are  such  that 
a  new  name  is  deemed  necessary.  The  jugal  canal  is,  I  believe,  a  new  formation  in 
Amphibia.  The  transverse  canal  of  the  amphibian  skull  is  homologous  with  the 
"occipital  cross-commissure." 

The  figure  (see  fig.  6)  is  a  composite  picture  of  the  lateral-line  system  of  the 
higher  or  truly  stegocephalous  Amphibia.  The  outline  of  the  skull  is  based  on  that 
of  Eryops.  All  of  the  canals  do  not  exist  on  any  one  skull  or  in  any  one  order,  but 
all  are  found  somewhere  in  the  group. 


Fig.  ii. 

A.  Skull  of  Eoserpeton  tenuicorne  Cope,  showing  arrangement  of  cranial  elements.    X  2.   fr,  frontal; 

j,  jugal;  mx,  maxilla;  n,  nasal;  or,  orbit;  par,  parietal;  pof,  postfrontal;  pmx,  premaxilla;  po,  post- 
orbital;  pp,  postparietal;  qj,  quadrat  ojugal ;  sq,  squamosal;  spt,  supratemporal;  tab,  tabularc. 

B.  Outline  of  skull  of  Ceraterpeton  galvani  Huxley  from  the  Carboniferous  of  England.   Heavy  broken 

lines  show  the  distribution  of  lateraldine  canals.  X  I.  (After  Andrews.)  fr,  frontal;  par,  parietal; 
or,  orbit;  po,  postorbital;  pp,  postparietal;  spt,  supratemporal;  tab,  tabularc. 

The  canals  have  been  described  in  all  known  orders  of  fossil  Amphibia  and  the 
system  is  found  likewise  in  all  the  living  orders,  including  the  Gymnophiona,  which 
have  "a  strong  line  of  lateral  sense-organs"  (Gadow).  In  the  Branchiosauria,  the 
earliest  of  the  true  Amphibia  (Euamphibia)  and  ancestral  to  the  modern  Caudata, 
the  lateral-line  system  is  known  on  the  tails  of  two  genera  (462,  478)  from  the  Mazon 
Creek,  Illinois,  shales — Micrerpeton  and  Eumicrerpeton.  The  system  as  there 
defined  has  been  fully  discussed  in  the  description  of  the  anatomical  details  of  the 
species,  to  which  reference  may  be  made  for  further  data  (pp.  52-60) .  Suffice  it  to  say 
here  that  the  system  of  sense-organs  there  preserved  is  identical  with  that  of  the 
larval  Necturus;  the  lines  arising  as  a  median  from  the  tip  of  the  tail  and  a  dorsal 
springing  from  the  median  at  a  distance  of  a  few  millimeters  from  the  tip  of  the  tail. 
The  lines  are  more  evident  on  account  of  the  fact  that  the  lateral-line  sense-organs 
were  located  under  specialized  pigmented  scales.    The  significance  of  the  close  simi- 


34  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

larity  between  the  arrangement  of  the  lateral-line  systems  on  the  tail  of  Necturus, 
Micrerpeton,  and  Eumicrerpeton  is  doubtless  of  genetic  (459)  importance,  indicating 
the  origin  of  the  caudate  Amphibia  from  the  Branchiosauria  by  a  degenerative  or 
recessive  evolution  in  other  structural  characters.  This  system  of  sense-organs  has 
been  described  in  no  other  branchiosaurian. 

The  Microsauria  (458)  are  exceedingly  interesting  in  possessing  a  very  peculiar 
type  of  lateral-line  system.  It  is  known  in  a  few  forms  and  in  one  specimen  espe- 
cially well  {Erpetosaurus  tabulatus)  (fig.  22,  G).  In  this  species,  which  is  represented 
by  a  single  imperfect  skull,  there  are  evidences  of  a  nearly  complete  lateral-line  sys- 
tem of  canals  and  pits.  The  occipital  cross-commissure  is  represented  on  the  pos- 
terior border  of  the  skull  by  a  row  of  elongate  pits  such  as  Andrews  described  for 
Ceraterpeton  (8).  I  fail  to  find  in  American  species  the  pores  described  by  Andrews. 
The  temporal  canal  forms  with  the  jugal  canal  a  complete  ring,  much  as  it  is  in  Tre- 
matosaurus,  only  in  Erpetosaurus  tabulatus  the  temporal  canal  does  not  touch  the 
tabulare.  I  think  there  are  indications  of  a  connection  of  the  temporal  canal  with 
the  supraorbital.  The  temporal  canal  cuts  the  supratemporal,  the  squamosal,  and 
jugal.  The  jugal  canal  lies  for  the  most  part  on  the  supratemporal  and  quadrato- 
jugal,  and  joins  the  infraorbital  on  the  jugal.  A  portion  only  of  the  infraorbital 
canal  is  preserved.  There  is  also  a  portion  of  the  supraorbital  canal.  It  seems  not 
to  be  connected  with  the  temporal  canal,  although  there  is  a  possible  indication  of 
this  connection.  The  supraorbital  crosses  the  frontal,  prefrontal,  and  a  part  of  the 
nasal.  The  squamosal  element  is  peculiar  in  Erpetosaurus  tabulatus  in  that  it  is 
excluded  from  the  parietal  by  the  extension  of  the  tabulare  and  postorbital.  This 
condition  is  found  in  several  other  species  of  the  Microsauria.  It  will  be  noticed 
that  with  the  changed  condition  of  the  squamosal  the  temporal  canal  has  changed 
also,  and  this  is  further  proof  of  the  close  connection  between  the  cranial  elements 
and  the  lateral-line  canals,  as  Allis  has  maintained  for  Amia.  (See  in  this  connection 
C.  J.  Herrick,  Journ.  Comp.  Neurol.,  vol.  11,  p.  224,  1901.) 

The  Diplocaulia,  an  amphibian  order  allied  to  the  Branchiosauria  (477)  and 
through  them  to  the  Caudata,  have  the  lateral-line  system  apparently  well-devel- 
oped. The  skulls  are  always  crushed  flat,  so  that  the  canals  are  nearly  obliterated. 
On  the  mandible,  however,  the  canals  are  clearly  distinct  and  apparently  run  the 
entire  course  around  the  mandibular  rami.  On  a  well-preserved  skull  of  Diplocaulus 
magnicornis  Cope  there  are  indications  of  three  lateral-line  canals  (477,  pi.  1).  The 
infraorbital  is  clearly  marked  as  a  well-defined  groove  just  below  the  orbit.  The 
supraorbital  is  indicated  only  for  a  short  distance,  and  there  are  indications  of  the 
temporal  canal.  The  operculo-mandibular  canal  has  its  course,  for  the  most  part, 
near  the  middle  of  the  rami,  but  as  it  approaches  the  posterior  angle  of  the  mandible 
it  suddenly  changes  its  course  and  drops  down  to  the  lower  edge,  only  to  rise  again 
and  to  come  out  strongly  marked  near  the  median  plane  on  the  posterior  angle  of 
the  mandible. 

The  Temnospondylia,  as  represented  by  Eryops,  Cricotus,  and  Archegosaurus, 
possess  well-developed  lateral-line  canals  (458).  H.  von  Meyer  (428)  many  years 
ago  made  out  the  course  of  the  canals  in  Archegosaurus.    The  greater  part  of  the  fol- 


MORPHOLOGY   OF   THE   COAL   MEASURES   AMPHIBIA.  35 

lowing  description  is  based  on  Eryops  megacephalus  Cope  from  the  Texas  Permian. 
The  entire  surface  of  the  cranial  elements  in  Eryops,  as  in  other  of  the  Stegocephala 
(458),  is  covered  with  coarse  pits.  The  fossae  are  present  even  in  the  bottoms  of  the 
grooves  which  represent  the  lateral-line  system,  and  are  more  marked  in  the  mem- 
bers of  the  Temnospondylia  than  in  the  Stereospondylia. 

The  occipital  cross-commissure  occurs  in  a  well-developed  form  in  Eryops.  It  is 
short  and  ends  abruptly  within  the  limits  of  the  tabulare.  Its  ends  are  occupied 
by  large  pits.  The  commissure,  as  in  Amia,  grooves  the  postparietal  and  the  tabu- 
lare elements.  There  is  no  evidence  of  an  anterior  commissure.  I  think  there  is 
evidence  of  a  temporal  canal  on  the  left  side  of  the  skull,  but  am  not  sure.  The 
jugal  and  infraorbital  canals  are  well  developed  and  strongly  connected.  The  jugal 
canal  starts  far  back  on  the  supratemporal,  and  after  curving  around  over  the  quad- 
ratojugal  joins  the  infraorbital,  or  rather  becomes  a  part  of  that  canal,  somewhere 
on  the  jugal.  There  is  nothing  unusual  in  the  infraorbital.  The  antorbital  commis- 
sure is  well  developed.  It  is  longer  and  better  developed  than  in  any  other  known 
form.  The  supraorbital  canal  starts  in  the  region  of  the  orbit,  and  after  curving 
downwards  to  meet  the  antorbital  commissure,  ends  abruptly  anterior  to  the  nostril. 
There  are  faint  traces  of  a  lateral-line  canal,  the  operculo-mandibular,  on  a  poorly 
preserved  mandible  of  Eryops.  It  does  not  differ  greatly  from  that  described  below 
for  Anaschisma. 

Although  Archegosaurus  has  been  known  for  more  than  a  century,  we  have  had 
no  adequate  discussion  of  the  manner  of  occurrence  of  the  lateral-line  canals.  Bur- 
meister  (80)  gave  a  figure  of  the  canals  as  he  thought  they  occurred  on  the  cranium, 
but  H.  von  Meyer  (428)  states  that  the  representation  is  inaccurate,  and  they  seem 
to  be  based  largely  on  Trematosaurus. 

The  lateral-line  canals  occur  in  well-developed  form  on  the  skulls  of  the  Stereo- 
spondylia. The  sutures  between  the  elements  of  the  skull  are  usually  clearly  marked 
by  smooth,  narrow  grooves.  The  lateral-line  canals  can  always  be  distinguished 
from  the  sutural  grooves  by  the  shape  of  the  bottom,  being  U-shaped  in  the  former 
and  V-shaped  in  the  latter.  The  lateral-line  canals  also  at  times  have  their  bottoms 
roughened  by  pits  occurring  in  them ;  the  sutural  grooves  always  have  smooth  bot- 
toms. The  lateral-line  canals  are  usually  rather  shallow  and  sometimes  broad, 
with  the  edges  of  the  grooves  more  or  less  perpendicular,  but  in  Metoposaurus  the 
canals  are  deep  and  the  borders  are  sharply  incised. 

The  temporal  canals  in  Anaschisma  from  the  Triassic  (49)  of  Wyoming  are  rep- 
resented by  broken  furrows.  The  portions  preserved  exhibit  the  usual  downward 
tendency  to  unite  with  the  infraorbital  on  the  postorbital  element.  In  its  course 
forward  from  the  epiotic  the  temporal  canal  cuts  the  squamosal.  The  supraorbital 
canal  has  an  unusually  deviating  course  in  Anaschisma,  but  aside  from  the  minor 
twists  and  curves  it  does  not  differ  essentially  from  the  same  canal  in  other  forms. 
It  ends  abruptly  on  the  anterior  end  of  the  muzzle.  In  its  course  it  gives  off  the  ves- 
tige of  an  antorbital  commissure  which  tends  to  join  a  vestige  from  the  infraorbital 
canal.  The  jugal  canal  begins  broadly  at  the  very  posterior  edge  of  the  skull  as 
though  it  were  continued,  as  it  undoubtedly  was,  to  the  body  of  the  animal.    In  its 


36  THE    COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

course  forward  it  joins  the  infraorbital  canal  on  the  jugal.  The  course  of  the  infra- 
orbital is  not  unusual  in  any  respect.  There  is  no  anterior  commissure  on  the  skull, 
nor  is  the  occipital  cross-commissure  developed  on  either  skull  of  the  genus  at  hand. 

There  are  distinct  evidences  of  an  operculo-mandibular  lateral-line  canal  on  the 
mandibles.  The  canal  enters  the  mandible  on  the  surangular  and  passes  forward 
around  the  mandible  as  described  for  Diplocaidus  (477). 

Other  members  of  the  Stereospondylia,  such  as  Mastodonsaurus,  Metoposaurus, 
and  Trematosaurus  possess  well-developed  lateral-line  canals,  but  the  above  descrip- 
tion fits,  in  a  general  way,  the  condition  in  all  genera;  and  for  our  present  purposes 
that  will  suffice. 


CHAPTER  V. 

THE  AMPHIBIA  OF  THE  DEVONIAN  AND  MISSISS1PPIAN  OF  NORTH  AMERICA. 

Evidences  of  the  earliest  land  vertebrates  are  exceedingly  scanty  in  the  strata 
between  the  close  of  the  Silurian  and  the  opening  of  the  Coal  Measures,  being  repre- 
sented solely  by  footprints.  In  the  Devonian  our  knowledge  of  the  group  is  con- 
fined to  a  single  footprint,  and  in  the  Mississippian  to  series  of  footprints  from 
several  localities.  These  have  been  described  by  Lea  (371),  Rogers  (Geology  of 
Pennsylvania,  pt.  11,  1856,  p.  831),  Barrell  (21),  Dawson  (223),  and  Branson  (50). 
The  last-named  author  has  described  a  new  species  from  the  Mississippian  of  Giles 
County,  Virginia.  His  description  of  the  footprints,  with  a  photograph  of  one  of 
the  series,  are  published  herewith  (plate  18,  fig.  3).  Branson  (50)  has  given  a  resume 
of  the  knowledge  of  Mississippian  Amphibia  in  North  America. 

Thinopus  antiquus  Marsh,  1896. 
MARSH,  Am.  Jour.  Science,  H,  p.  374,  Nov.  1896,  with  figure. 


Type:    Specimen  No.  784,  Yale  University  Museum. 
Horizon :    Near  top  of  Chemung,  in  the  upper  Devonian. 


[The]  "specimen  shows  one  vertebrate  footprint  in  fair  preservation,  and  with  it  part 
of  another  of  the  same  series.  These  impressions  are  of  much  interest,  both  on  account  of 
their  geological  age  and  the  size  and  character  of  the  footprints  themselves.  The  one  best 
preserved  [fig.  12]  is  nearly  4  inches  in  length,  2.25  inches  in  width,  and  was  apparently 
made  by  the  left  hind  foot.  On  the  inner  side  in  front 
of  the  heel,  a  portion  of  the  margin  is  split  off,  and 
this  may  have  contained  the  imprint  of  another  toe. 
The  other  footprint  was  a  short  distance  in  front,  but 
only  the  posterior  portion  is  now  preserved  in  the 
present  specimen.  It  is  probably  the  imprint  of  the 
forefoot. 

"The  specimen  [plate  18,  fig.  4]  .  .  .  was   .  .  . 
found  in  the  town  of  Pleasant,  one  mile  south  of  the 
Allegheny  River,  Warren  County,  Pennsylvania,  by 
Dr.  Charles  E.  Beecher,  who  presented  it  to  Yale  Fie  12-Copy  of  Mush's  drawing  of  footprint  of 
rv  ,  •       ,-,,  •  Thinopus  antiquus,  from  the  Devonian  of  Penn- 

Museum,  where  it  still  remains.  sylvania.   X  'A. 

"The  geological  horizon  is  near  the  top  of  the 
Chemung  in  the  upper  Devonian.    In  the  same  beds  are  ripple  marks,  mud  cracks,  and 
impressions  of  rain  drops,  indicating  shallow  water  and  shore  deposits.     Land  plants  are 
found  in  the  same  general  horizon.     Marine  molluscs  also  occur,  and  one  characteristic 
form  (Nuculana)  is  preserved  in  the  footprint  slab"  (Marsh). 

This  still  remains  after  nearly  20  years  the  only  evidence  of  air-breathing  verte- 
brates in  the  Devonian  of  the  world. 

Dromopus  ad  uncus  Branson. 
Branson,  Jour.  Geol.,  .win,  No.  4,  pp.  356-358,  fig.  '.  «910- 

Type  and  other  specimens  in  Oberlin  College  Museum. 

Horizon  and  type  locality:   Near  the  bottom  of  the  Hinton  formation  in  Giles 

County,  Virginia.     (Plate  18,  fig.  3.) 

37 


38  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

The  following  description  of  the  shales  and  footprints  are  from  Dr.  Branson's 
paper  cited  above: 

"The  Hinton  shales,  like  the  Mauch  Chunk,  seem  to  have  been  subaerial  in  origin  and 
are  made  up  for  the  most  part  of  variegated  shales  interbedded  with  thin  layers  of  argil- 
laceous, fine-grained  sandstone.  The  footprints  occur  in  fine-grained  sandstone,  and  remains 
of  land  plants  are  not  uncommon  in  the  same  beds. 

' '  Twenty- two  footprints  made  by  one  animal  walking  in  a  straight  course  were  collected 
in  a  slab.  They  give  the  impression  of  having  been  made  by  a  bipedal  animal  for  part  of 
the  distance,  but  after  the  fourth  print  of  the  right  foot  impressions  of  the  forefeet  appear. 
The  hindfeet  had  5  digits,  the  middle  digit  being  longest  and  the  2  inside  of  it  being  only 
slightly  shorter  and  lying  close  together.  Their  outer  ends  were  slender  and  flexible  and 
usually  curved  inward  toward  the  middle  toe.  The  2  outer  digits  formed  wide  angles  with 
the  middle  one  and  were  shorter  than  the  inner  ones.  The  second  toe  was  webbed  to  within 
8  mm.  of  the  tip,  the  third  toe  to  within  23  mm.  of  the  tip.  The  impression  of  the  web  is 
well  preserved  in  only  one  impression  of  the  hindfoot. 

"The  forefeet  had  4  digits.  The  3  inner  digits  were  subequal  in  length,  the  2  inner 
being  more  flexible  and  incurved  near  the  ends.  The  outer  digit  is  two- thirds  as  long  as 
the  second.  The  webbing  extends  about  half  the  length  of  the  digits.  The  heel  impression 
is  broader  than  that  of  the  hindfoot. 

"Measurements  of  Dromopus  aduncus  Branson. 

Tip  of  toe  to  tip  of  toe  in  first  prints 21  cm. 

After  appearance  of  forefeet  the  impressions  are  the  following  distances  apart :  165  mm.,  40  mm., 
85  mm.,  70  mm.,  80  mm.,  40  mm.,  150  mm.,  then  back  to  20  and  21  cm. 

mm. 

Length  of  hindfeet 60 

Width  of  hindfeet 20  to  25 

Length  of  forefeet 45" 


CHAPTER  VI. 

A    HISTORY    OF    THE    CLASSIFICATION    OF     THE     AMPHIBIA,     WITH    ESPECIAL 
REFERENCE  TO  THE  SPECIES  FROM  THE  COAL  MEASURES. 

It  has  been  necessary,  in  the  course  of  the  present  study,  to  review  thoroughly 
the  classifications  which  have  been  proposed  for  the  group.  A  classification  of  some 
sort  is  necessary  for  the  proper  grouping  of  the  species  which  have  been  recovered 
from  the  Coal  Measures  deposits  of  this  continent,  and  my  reason  for  publishing 
this  relatively  dry  material  is  that  the  classifications  formerly  proposed  (469),  as 
well  as  the  one  here  given,  may  have  a  proper  historical  background. 

The  review  of  the  proposed  systems  of  classification  has  been  much  facilitated 
by  the  discovery,  in  the  University  of  Chicago,  of  some  notes  by  the  late  Dr.  George 
Baur  on  the  "  Stegocephali."  The  notes  were  not  discovered  until  after  the  literature 
had  been  pretty  thoroughly  covered,  and  it  was  a  source  of  some  gratification,  on 
comparing  notes  with  those  of  Dr.  Baur,  to  find  but  few  omissions.  Whether  Dr. 
Baur  had  ever  contemplated  a  work  on  the  Stegocephala  or  not  I  have  been  unable 
to  learn,  but  it  is  certain  that  he  carefully  and  laboriously  went  through  the  litera- 
ture on  the  subject  and  copied  by  hand  the  classifications  of  each  author  from  1842 
to  1895,  together  with  other  notes  of  interest  on  the  structure,  distribution,  and 
phylogeny,  including  many  tracings.  The  classifications  given  below  are  taken,  in 
part,  from  his  notes,  although  all  references  have  been  verified  with  the  original 
sources. 

The  first  attempt  to  combine  in  classification  the  knowledge  of  the  extinct  and 
recent  amphibians  was  made  by  Johannes  Jacob  von  Tschudi  in  1839  (574).  Pre- 
vious to  that  time  Goldfuss  (295)  and  von  Meyer  (418)  had  described  various 
species  of  salamanders  and  frogs  from  the  Tertiary  deposits  of  Switzerland,  and 
these  Tschudi  considered  in  his  following  classification: 

A.  Ranae.  B.  Cceciliae. 

a.  Hylcc.  a.  Cacilue. 

b.  Cystignatlu.  C.  Salamandrinae. 

c.  Ranee.  a.  Pleurodcles. 

d.  Ceratophrydes.  b.  Salamatidrce. 
c.  Bombinatores.  c.  Tritones. 

f.  Bufones.  d.  Tritonides. 

g.  PipcB.  D.  Protoideae. 

Although  the  remains  of  Mastodonsaurus  had  been  known  and  widely  com- 
mented on  for  several  years  before  Tschudi  proposed  this  scheme,  he  does  not 
include  this  genus  in  his  classification  of  the  Amphibia,  for  the  reason  that  for 
nearly  a  quarter  of  a  century  after  the  discovery  of  the  labyrinthodonts  they  were 
regarded  as  reptiles,  even  so  eminent  an  authority  as  von  Meyer  (423)  including 
them  in  his  "System  der  fossilen  Saurier."    The  view  that  the  labyrinthodonts  were 

39 


40  THE   COAL  MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

reptiles  was  at  times  disputed,  but  no  one  seemed  to  pay  any  attention  to  the  argu- 
ment of  Quenstedt  in  1850  that  "Die  Mastodonsaurier  im  grunen  Keupersand- 
stein  Wurtemburgs  sind  Batrachier"  (527),  nor  to  the  contention  of  Vogt  (581)  in 
1854  that  "  Archegosaurus  und  alle  Labyrinthodonten  sind  Amphibien,  nicht  Rep- 
tilien." 

In  1842  von  Meyer  (420)  proposed  to  include  all  the  early  forms  allied  to  the 
Mastodonsaurus  in  the  " Labyrinthodontes. "    His  definition  of  the  group  follows: 

Labyrinthodontes  :  Saurier  deren  Zahn-Struktur  jener  ahnlich  ist,  welche  in  den  nach  prismat- 
ischer  Art  gebauten  Saugethier-Zahnen  wahrgenommen  wird,  u.s.w. 

I.  Mastodonsaurus  Jaeger.     (Salamandroides  Jaeger,  Batrachosaurus  Fitzinger,  Labyrinth- 

odon  Owen.)    M.  Jaegeri  Meyer. 
II.  Capitosaurus  Miinster. 
C.  arenaceus  Miinst. 
C.  robustus  Meyer. 
III.  Metopias  Meyer. 

M.  diagnosticus  Meyer. 

Three  years  later  von  Meyer  (423)  proposed  his  "System  der  fossilen  Saurier," 
where  the  extinct  Amphibia  are  treated  as  follows: 

Labyrinthodontes. 

1.  Prosthopthalmi    (Augen-hohlen    in    der    vordern    Halfte  der    Schadel-Lange)    Metopias 

Meyer Keuper. 

2.  Mesopthalmi  (Augen-hohlen  in  der  mitte  der  Schadel) 

Mastodonsaurus  Jaeger — Keuper,  Muschelkalk. 

3.  Opisthopthalmi    (Augen-hohlen   in   der   hintern   Halfte   der  Schadel-Lange)  Capitosaurus 

Miinster Keuper. 

4.  Labyrinthodonten  ungewisser  Stellung. 

Labyrinthodon   Owen Keuper. 

Xestorrhytias  Meyer Muschelkalk. 

Odontosaurus  Meyer Bunter  Sandstein. 

Trematosaurus  Braun Bunter  Sandstein. 

No  other  classification  was  proposed  for  the  extinct  Amphibia  for  15  years,  when 
Owen  (512)  in  1859  proposed  the  new  order  Ganocephala  and  retained  von  Meyer's 
Labyrinthodontes  under  Labyrinthodontia.    Owen's  classification  is  as  follows: 

Class — Reptilia. 
Order    I.  Ganocephala. 

Genera:  Archegosaurus,  Dendrerpeton,  Raniceps. 
Order  II.  Labyrinthodontia. 

Genera:   Mastodonsaurus,   Anisopus,    Trematosaurus,   Metopias,    Capitosaurus,    Zygosaurus, 
Xestorrhytias. 

In  his  Paleontology  published  in  1861,  Owen  gives  the  same  classification,  but 
adds  new  genera. 

Huxley  in  1863  (332)  did  not  accept  Owen's  Ganocephala,  but  instead  proposed 
the  following: 

Labyrinthodontia. 

A.  Archegosauria.    Archegosaurus,  Pholidogaster . 

B.  Mastodonsauria.    Mastodonsaurus,  Labyrinthodon,  Capitosaurus,  Trematosaurus. 

In  the  same  year  Dawson  proposed  (208)  the  term  Microsauria  to  include  the 
genera  Hylonomus,  Dendrerpeton,  and  Hylerpeton,  all  known  from  the  Carboniferous 


HISTORY  OF   CLASSIFICATION   OF  AMPHIBIA.  4 1 

rocks  of  Nova  Scotia.  Two  years  later  Cope  proposed  the  new  order  Xenorhachia 
(105)  for  the  reception  of  the  form  Amphibamus  grandiceps  from  the  Coal  Measures 
of  Illinois.  He  gave  as  the  characters  of  this  order  cartilaginous  vertebras  and  the 
absence  of  ribs. 

In  1866  Owen  proposed  (516)  the  most  elaborate  and  comprehensive  scheme  of 
classification  which  had  thus  far  been  offered.    His  classification  is  as  follows: 

Subclass  Dipnoa. 

Order  Ganocephala  (extinct).    Genera:  Archegosaurus,  Dendrerpeton,  etc. 
Order  Labyrinthodontia.    Genera:  Labyrinthodon,  Rhombopholis,  etc. 
Order  Batrachia. 

Suborder  Ophiomorpha.    Family:  Ccecilidae. 
Suborder  Ichthyomorpha.     Family:  Proteidae,  Salamandridae. 
Suborder  Theriomorpha  (Anura). 
Family  1.  Aglossa. 
Family  2.  Ranida. 
Family  3.  Hylidce, 
Family  4.  Bufonidce. 

Haeckel  the  same  year  proposed  (312)  an  entirely  different  scheme  of  classifica- 
tion and  in  some  respects  more  acceptable  than  Owen's.  Haeckel's  classification 
is  as  follows : 

Class  Amphibia. 

Subclass  I.  Phractamphibia. 
Ordnung  1.  Ganocephala. 

Genera:  Archegosaurus,  Dendrerpeton,  Raniceps. 
Ordnung  2.  Labyrinthodonta. 

Genera:  Baphetes,  Zygosaurus,  Mastodonsaurus,  Trematosaurus,  Capitosaurus. 
Ordnung  3.  Peromela. 
Subclass  II.  Lissamphibia. 
Ordnung  1.  Socobranchia. 

Genera:  Siren,  Proteus,  Menobranchus,  etc. 
Ordnung  2.  Sozura  (Caudata). 

Genera:  Cryptobranchus,  Triton,  Salatnandra. 
Ordnung  3.  Anura  (Ecaudata). 

Families:  Aglossa,  Bufonidce,  Ranidce. 

This  classification  is  further  elaborated  in  the  edition  of  1895. 
Cope  in  1868  proposed  (no)  the  scheme  of  classification  which  was  in  use  for 
some  time,  although  it  has  since  suffered  some  change.    His  classification  follows: 

Batrachia. 

Order  1.  Trachystoma. 
Order  2.  Proteida. 
Order  3.  Urodela. 
Order  4.  Gymnophiona. 
Order  5.  Stegocephali. 

Suborder  Xenorhachia. 

Amphibamus  grandiceps  Cope. 
Suborder  Microsauria. 

Genera:  Pelion  Wyman,  Hylonomus  Dawson,  Pariostegus  Cope,  Dendrerpeton 
Owen,    Hylerpeton    Owen,    Brachydectes    Cope,    Sauropleura    Cope,    CEsto- 
cephalus  Cope,  Molgophis  Cope. 
Suborder  Labyrinthodontia. 

Genera:  Dictyocephalus  Leidy,  Centemodon  Lea,  Baphetes  Owen,  Eupelor  Cope. 


42  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

Huxley  in  1869  proposed  (335)  the  following  classification,  which  does  not  differ 

essentially  from  that  proposed  in  1863: 

Amphibia. 

Order  1.  Urodcla. 
Order  2.  Batrachia. 
Order  3.  Gymnophiona. 
Order  4.  Labyrinthodontia. 

Suborder  Archegosauria. 

Suborder  Mastodonsauria. 

The  next  classification  of  the  extinct  Amphibia  of  any  importance  was  that 
devised  by  the  committee  (450)  for  the  British  Association  in  1874.  This  committee 
was  formed  of  Huxley,  Harkness,  Henry  Woodward,  Thompson,  and  Brigg,  with 
Miall  as  secretary.  This  classification  is,  however,  too  cumbersome  and  has  never 
come  into  general  use,  and  indeed  none  but  English  authors  have  paid  it  a  great  deal  of 
attention,  although  the  contribution  was  a  valuable  one.  The  group  Aistopoda,  which 
was  the  ninth  group  proposed  by  the  committee,  has  generally  been  accepted  as  the 
group  represented  by  the  snake-like  forms.     The  committee's  classification  follows: 

Labyrinthodontia. 

Section  I.  Euglypta. 

Genera:  Mastodonsaurus,  Jaeger;  Capitosaurus,   Miinst. ;  Pachygonia,  Huxley;  Tremato- 
saurus,  Braun;  Gonioglyptus,  Huxley;  Metopias,  von  Meyer;  Iuibyrintliodon,  Owen; 
Diadetognathus,  Miall;  Dasyceps,  Huxley;  Anthracosaurus,  Huxley. 
Section  II.  Brachyopina. 

Genera:  Brachyops,   Owen;   Micropholis,    Huxley;   Rhinosaurus,    Waldheim;    Bothriceps, 
Huxley. 
Section  III.  Chauliodonta. 

Genera:  Loxomma,  Huxley;  Zygosaurus,  Eich.;  Melosaurus,  Meyer. 
Section  IV.  Arthroodonta. 

Genera:  Batrachiderpeton,  Hancock  and  Atthey;  Pteroplax,  Hancock  and  Atthey. 
Section  V.  An  uncharacterized  group. 

Genera:  Pholidogaster,  Huxley;  Ichthyerpelon,  Huxley;  Pholiderpelon,  Huxley. 
Section  VI.  Archegosauria,  von  Meyer. 

Genera:  Archegosaurus,  Goldfuss. 
Section  VII.  Heleothrepta. 

Genera:  Lepterpeton,  Huxley. 
Section  VIII.  Nectridea. 

Genera:  Urocordylus,  Huxley;  Keraterpeton ,  Huxley. 
Section  IX.  Aistopoda. 

Genera:  Ophiderpeton,  Huxley;  Dolichosoma,  Huxley. 
Section  X.  Microsauria,  Dawson. 

Genera:  Dendrerpeton,  Owen;  Hylonomus,  Dawson;  Hylerpeton,  Owen. 

The  schemes  of  classification  used  for  the  next  10  years  did  not  depart  in  any 
appreciable  degree  from  those  already  given. 

In  1875  Cope,  in  his  Check-list  of  North  American  Batrachia  and  Reptilia  (120), 

with  a  systematic  list  of  the  higher  groups,  published  the  following  classification  as 

being  "adopted  provisionally  by  the  Smithsonian  Institution": 

Class  Batrachia. 

Order  Anura.    (Anura,  Dumeril;  Salientia,  Merrem,  Gray.) 

Raniformia.    (Raniformia,  Cope,  Nat.  Hist.  Rev.,  v,  114,  1865.) 

Families:  Ranidce,  Colostheidce. 
Firmisternia.     (Bufonoid  Raniformia,  Cope,  Jour.  Acad.  Nat.  Sci.,  Philadelphia,  n.s.,  vi, 
190,  1867.) 
Families:  Dendrobatidee,  Phryniscidce,  EngystomidcB,  Breviceptidce. 


HISTORY  OF  CLASSIFICATION  OF  AMPHIBIA.  43 

Class  Batrachia — Continued. 
Order  Anura — Continued. 

Gastrechmia.    (Gastrechmia,  Cope,  Jour.  Acad.  Nat.  Sci.,  Philadelphia.,  n.s.,  vi,  198,  1867.) 

Family:  Hemisidce. 
Bufoniformia.    (Bufoniformia,  Dumeril  et  Bibron,  partim;  Cope,  partim.) 

Families:  Rhinophrynidw,  Bufonida,  Batrachophrynidw. 
Aglossa. 

Family:  Pipidoe. 
Odontaglossa. 

Family:  Dactylethrida. 
Arcifera.    (Arcifera,  Cope,  N.  H.  Rev.,  v,  104,  1865.) 

Families:  Cystignathidce,   Hemiphractidce,   Hylidce,   Scaphiopidce,   Pelodytida,  Astero- 
phrydidce,  Discoglossidee. 
Order  Stegocephali.    (Stegocephali  Cope,  Proc.  Acad.  Nat.  Sci.,  Philadelphia,  1868,  209.) 
Labyrinthodontia . 

Families:  Baphetidce  Cope,  Anthracosauridw  Cope. 
Ganocephala. 

Family :  Colosteidce  Cope. 
Microsauria. 

Families:  Phlegethontiidce  Cope,  Molgophidw  Cope,  Ptyoniida;  Cope,  Tuditanidx  Cope, 
Peliontidce  Cope. 
Order  Gymnophiona.    (Gymnophiona  Muller.) 

Family:  Caciliida  Gray,  1850. 
Order  Urodela. 

Families:  Pleurodelidce  Gray,  1858;  Salamandridw  Gray,  1858;  Hynobiidce  Cope,  1866; 
Desmognathidce  Cope,  1866;  Thoriidm  Cope,  1869;  Plethodontidce  Cope, 
1866;  Amblystomidee    Cope,    1866;  Menopomidm   (Protonopsidce   Gray, 
1850),  Amphiumidm  Cope,  1866;  Cocytinidce  Cope. 
Order  Proteida. 

Family:  Proteida  Gray,  1850. 
Order  Trachystomata. 

Family:  Sirenidw  Gray,  1850. 

In  1885  Cope  proposed  2  new  orders,  which  he  arranges  with  the  other  orders 

already  known,  as  follows: 

Order  I.  Rachitomi.  Order      V.  Urodela. 

Order  II.  Embolomeri.  Order    VI.  Trachystomata. 

Order  III.  Stegocephali.  Order  VII.  Anura. 

Order  IV.  Proteida. 


Batrachia. 


The  new  order  Rachitomi  was  to  include  forms  like  Eryops  and  the  new  order 
Embolomeri  was  to  include  forms  like  Cricotus;  the  other  orders  were  as  they  had 
been  given  before. 

Zittel  in  1888  proposed  (642)  the  next  classification  of  any  note  in  his  "Hand- 

buch  der  Paleontologie,"  where  it  stands  as  follows: 

Classe  Amphibia. 

Ordnung  1.  Stegocephali.  Ordnung  3.  Urodela. 

Unterordnung  1.  Lepospondyli.  Unterordnung  1.  Ichthyoidea. 

Familie  1.  Branchiosauridce  Fritsch.  Familie  1.  Phanerobraiwlua. 

2.  Microsauria  Dawson.  "      2.  Cryptol>ra>ulihi. 

"       3.  Aistopoda  Miall.  Unterordnung  2.  Salamandrina. 

Unterordnung  2.  Temnospondyli.  Ordnung  4.  Anura. 

Genera:    Archegosaurus,  Eryops,  etc.  Unterordnung  r.  Phaneroglossa. 

Unterordnung  3.  Stereospondyli.  Familie  1.  Ranidw. 

Familie  1.  Gastrolepidoti.  "       2.  Bufonida. 

2.  Labyrinthodonta.  "      3-  Cystignathida  Cope. 

Ordnung  2.  Cceciliae.  "      4-  PelobaXidm  Boul. 

"      5.  Discoglossidee  Cope. 
"       6.  Palaobatraihidce  Cope. 


44  THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 

Lydekker  (393)  in  the  next  year  proposed  a  system  of  classification  which  did 
not  depart  widely  from  that  proposed  (450)  by  the  committee  of  the  British  Associ- 
ation for  1874.     Lydekker's  classification  is  as  follows: 

Class  Amphibia. 

Order  I.  Labyrinthodontia.  Order  II.  Apoda. 

Suborder  1.  Branchiosauria.  Order  III.  Caudata. 

Family  Protritonidce.  Family  Hylceobatrachid.ee. 

"      Apateonidce .  "      Sirenidw. 

Suborder  2.  Aistopoda.  "      Proteidoe. 

Family  Dolichosamatidce .  "      Amphiumidce. 

Suborder  3.  Microsauria.  "      Salamandridce. 

Family  Urocordylidce.  Order  IV.  Ecaudata. 

"       Limner petidce.  Family  Discoglossidce. 

"      Hyloplesionidce.  "       Pelobatidce. 

"       Microbrachidcz.  "      Palceobatrachidce. 

Suborder  4.  Labyrinthodontia  vera.  "      Cystignathidce. 

Family  Archegosauridce.  "      Ranidce. 

"      Diplospondylidce. 
"      Nyraniidce. 
"      Dendrer petidce. 
"       Anthracosauridce . 
"      Mastodonsauridce. 
Uncertain  family,  Eosaurus. 

In  1890  Doederlein  proposed  a  scheme  of  classification  which  is  notable  on 
account  of  the  peculiar  relations  which  it  expresses  between  the  groups — relations 
which,  in  reality,  do  not  exist.    His  classification  is  as  follows: 

Class  Amphibia. 

Ordnung   I.  Stegocephali. 

A.  Microsauria. 

Unterordnung  1.  Branchiosauri. 

Genera:  Branchiosaurus,  Dawsonia,  Melanerpeton,  Pelosaurus. 
Unterordnung  2.  Sauromorphi. 
Familie  1.  Hylonomidce. 
"     2.  Nectridce. 
"     3.  Aistopodidce. 

B.  Ganocephala. 

Unterordnung  1.  Rhachitomi.  * 
2.  Embolomeri. 
"  3.  Labyrinthodontia. 

Ordnung    II.  Urodela. 
Ordnung  III.  Gymnophiona. 
Ordnung  IV.  Anura. 

In  1890  Lydekker  used  the  same  classification,  with  minor  changes,  which  he  had 
used  in  his  Paleontology.  Credner,  who  wrote  at  about  the  same  time  (193),  fol- 
lowed Zittel's  classification.  Zittel  in  1895  merely  repeated  his  former  classification. 
In  1898  appeared  Smith  Woodward's  Paleontology,  where  the  following  scheme  is 
adopted : 

Class  Batrachia. 

Order     I.  Stegocephalia.  Order    II.  Gymnophiona. 

Suborder  1.  Branchiosauria.  Order  III.  Caudata. 

2.  Aistopoda.  Order  IV.  Ecaudata. 

"       3.  Microsauria. 

4.  Labyrinthodontia. 


HISTORY   OF   CLASSIFICATION   OF  AMPHIBIA.  45 

Hay's  Catalogue  of  Fossil  Vertebrata  of  North  America  contains  the  next  scheme 
for  the  classification  of  the  Amphibia  which  pays  especial  attention  to  the  extinct 
forms.    His  classification  is  as  follows: 

Class  Batrachia  Macartney,  1802. 
Order  Stegocephali  Cope,  1868. 

Suborder  Microsauria  Dawson,  1863. 
Family  Protrilonidm  Lydekker,  1889. 

Genera:  Amphibamus  Cope,  Pelion  Wym. 
Family  Molgophidw  Cope,  1875. 

Genera:  Phlegethontia  Cope,  Molgophis  Cope. 
Family  Hylonomida  Fritsch,  1883. 

Genera:  Hylonomus  Dawson,  Smilerpeton  Dawson,  Hylcrpeton  Owen,  Fritschia 
Dawson;  Brachydectes  Cope. 
Family  Ptyoniida  Cope,  1875. 

Genera:  Keraterpeton  Huxley,  Gistocephalus  Cope,  Piyonius  Cope,  Cienerpeton 
Cope. 
Family  Tuditanida  Cope,  1875. 

Genera:  Tudilanus  Cope,  Cocytinus  Cope. 
Family  Diplocaulidce  Cope,  1881. 
Genus:  Diplocaulus  Cope. 

Lepospondylous  Genera  of  uncertain  position:  Amblyodon  Dawson,  Hyphasma 
Cope,   Eurythorax   Cope,     Thyrsidium    Cope,    Pleuroptyx   Cope,    Cercario- 
morphus  Cope. 
Suborder  Apoecospondyli  Hay,  1902. 

Family  Dendrerpetontidce  Fritsch,  1889. 

Genera:  Dendrerpeton  Owen,  Baphetes  Owen,  Platystegos  Dawson. 
Family  Sauropleuridce  Hay,  1902. 

Genera:  Sauropleuta  Cope,  Leptophr actus  Cope. 
Family  Archegosauridce. 

Genera:  Trimerorhachis  Cope,  Dissoropkus  Cope. 
Family  Cricotidce  Cope,  1884. 

Genera:  Cricotus  Cope. 
Family  Anthracosaurida. 

Genus:  Eosaurus  Marsh. 
Family  Eryopidce  Cope,  1882. 

Genera:  Eryops  Cope,  Ichthycanthus  Cope,  Zatrachys  Cope,  Anisodexis  Cope, 
Acheloma  Cope. 
Family  Mastodonsaurida  Huxley,  1863. 

Genera :  Mastodonsaurus  Jaeger,  Eupelor  Cope,  Pariostegus  Cope,  Dictyocephalus 
Leidy. 
Order  Urodela. 

Genera:    Scapherpelon  Cope,  Hemitrypus  Cope. 
Order  Salientia  Laurcnti,  1768. 
Family  Ranidce. 

Genera:  Rana  Linne,  Eobatrachus  Marsh. 

This  classification  given  by  Hay  is  only  for  the  forms  which  occur  in  North 
America. 


CHAPTER  VII. 

CLASSIFICATION   OF  AMPHIBIA   ADOPTED   IN   THIS  WORK,  AND   A   LIST   OF   THE 
COAL  MEASURES  AMPHIBIA  FROM  NORTH  AMERICA. 

A  few  words  of  explanation  will  be  necessary  for  an  understanding  of  the  follow- 
ing classification.  The  term  Amphibia  Linne,  1758,  is,  according  to  Stejneger  (550), 
the  correct  term  for  the  entire  group,  and  this  term  is  adopted.  The  term  Stego- 
cephala,  formerly  used  as  a  group  name  for  the  entire  Carboniferous,  Permian,  and 
Triassic  Amphibia,  regardless  of  structure,  has  been  retained  as  a  third  subclass. 
The  choice,  so  far  as  priority  is  concerned,  between  Labyrinthodontia  and  Stego- 
cephala,  is  not  easy.  The  terms,  however,  imply  different  structures.  The  laby- 
rinthodonts  proper  have  stereospondylous  vertebrae,  while  the  Stegocephala  have 
either  temnospondylous  or  stereospondylous  vertebrae;  so  the  latter  term  has  been 
adopted. 

The  ordinal  terms  are  those  which  have  been  used  previously  as  subordinal, 
sectional,  or  family  names,  with  the  exception  of  the  new  ordinal  term  "Diplocau- 
lia"  (477).  The  term  Branchiosauria  is  well-established,  and  it  is  here  retained  with 
the  definitions  previously  given.  The  same  may  be  said  for  the  Microsauria, 
although  Dawson  first  (208)  regarded  it  as  a  family,  though  giving  the  term  an  ordi- 
nal form.  The  Aistopoda  are  not  entitled  to  consideration  as  a  group  for  reasons 
which  are  given  subsequently.  The  Temnospondylia  and  the  Stereospondylia,  the 
Embolomeri  and  the  Rachitomi  may  or  may  not  be  good  group  names,  but  they  have 
priority,  so  far  as  our  knowledge  of  structure  goes.  They  have  been  retained  in 
their  original  meanings.  They  have  been  variously  regarded  as  sections,  superf ami- 
lies,  groups,  and  subfamilies. 

The  following  list  of  species  is  arranged  according  to  the  proposed  scheme  of 
classification : 

Class  Amphibia  Linne,  1758.    Devonian  to  Recent. 

Subclass  Euamphibia  Moodie,  1909.    Coal  Measures  to  Recent. 

Order  Branchiosauria  Lydekker,  1889.    Coal  Measures  to  Permian. 
Family  Branchiosauridce  Fritsch,  1879. 

Micrerpeton  caudatum  Moodie,  Mazon  Creek. 
Eumicrerpeton  parvum  Moodie,  Mazon  Creek. 
Mazonerpeton  longicaudatum  Moodie,  Mazon  Creek. 
Mazonerpeton  costatum  Moodie,  Mazon  Creek. 
(?)  Sparodus  sp.  indet.    Dawson,  Nova  Scotia. 
Order  Caudata  Dumeril,  1806.    Coal  Measures  to  Recent. 

Suborder  Proteida  Cope,  1868.    Coal  Measures,  Eocene,  and  Recent. 
Family  Cocytinidoe  Cope,  1875. 

Cocytinus  gyrinoides  Cope,  Linton,  Ohio. 
Erierpeton  branchialis  Moodie,  Mazon  Creek. 
Hyphasma  laevis  Cope,  Linton,  Ohio. 
Order  Diplocaulia  Moodie,  191 2.    Coal  Measures  to  Permian. 
Family  Diplocaulidce  Cope,  1881. 

Diplocauhts  salamandroides  Cope,  Salt  Fork,  Illinois. 
(?)  Order  Salientia  Laurenti,  1768.    Coal  Measures  (?)  to  Recent. 
Family  Peliontida  Cope,  1875. 

Pelion  lyelli  Wyman,  Linton,  Ohio. 

46 


CLASSIFICATION   OF  AMPHIBIA   AND  LIST  OF  COAL  MEASURES  SPECIES.  47 

Class  Amphibia — Continued. 

Subclass  Lepospondylia  Zittel,  1887.    Coal  Measures. 
Order  Microsauria  Dawson,  1863.    Coal  Measures. 
Family  Hylonomidce  Fritsch,  1883. 

Hylanomus  latidens  Dawson,  Nova  Scotia. 

Hylonomus  lyelli  Dawson,  Nova  Scotia. 

Hylonomus  multidens  Dawson,  Nova  Scotia. 

Hylonomus  wymani  Dawson,  Nova  Scotia. 

Smilerpeton  aciedentatum  Dawson,  Nova  Scotia. 

Hylerpeton  dawsonii  Owen,  Nova  Scotia. 

Hylerpeton  intermedium  Dawson,  Nova  Scotia. 

Hylerpeton  longidentatum  Dawson,  Nova  Scotia. 

Fritschia  curtidentata  Dawson,  Nova  Scotia. 

(f)  Amblyodon  problematicum  Dawson,  Nova  Scotia. 
Family  Tuditanidce  Cope,  1875. 

Tuditanus  punctulatus  Cope,  Linton,  Ohio. 

Tuditanus  brevirostris  Cope,  Linton,  Ohio. 

Tuditanus  minimus  Moodie,  Cannelton,  Pa. 

Tuditanus  longipes  Cope,  Linton,  Ohio. 

Tuditanus  walcotti  Moodie,  Linton,  Ohio. 

Erpetosaurus  acutirostris  Moodie,  Linton,  Ohio. 

Erpetosaurus  minutus  Moodie,  Cannelton,  Pa. 

Erpetosaurus  obtusus  Cope,  Linton,  Ohio. 

Erpetosaurus  radiatus  Cope,  Linton,  Ohio. 

Erpetosaurus  sculptilts  Moodie,  Cannelton,  Pa. 

Erpetosaurus  tabulatus  Cope,  Linton.  Ohio. 

Erpetosaurus  tuberculatus  Moodie,  Linton,  Ohio. 

Odonterpeton  triangularis  Moodie,  Linton,  Ohio. 
Family  Stegopidw  Moodie,  1009. 

Stegops  divaricata  Cope,  Linton,  Ohio. 
Family  Urocordylidce  Lydekkcr,  1890. 

Diceratosaurus  punctolineatus  Cope,  Linton,  Ohio. 

Diceratosaurus  IcBvis  Moodie,  Linton,  Ohio. 

Diceratosaurus  robustus  Moodie,  Linton,  Ohio. 

Eoserpeton  tenuicorne  Cope,  Linton,  Ohio. 
Family  Amphibamida  Moodie. 

Amphibamus  grandiceps  Cope,  Mazon  Creek. 

Amphibamus  thoracatus  Moodie,  Mazon  Creek. 

Cephalerpeton  ventriarmatum  Moodie,  Mazon  Creek. 
Family  Nyraniidce  Lydekker,  1889. 

Ichthyerpcton  squamosum  Moodie.    Linton,  Ohio. 

Cercariomorphus  parvisquamis  Cope,  Linton,  Ohio. 
Family  Ptyoniida:  Cope,  1875. 

Ptyonius  nummifer  Cope,  Linton,  Ohio. 

Ptyonius  marshii  Cope,  Linton,  Ohio. 

Ptyonius  vinchellianus  Cope,  Linton,  Ohio. 

Ptyonius  pectinatus  Cope,  Linton,  Ohio. 

Ptyonius  serrula  Cope,  Linton,  Ohio. 

CEstocephalus  remex  Cope,  Linton,  Ohio. 

Qistocephalus  reciidens  Cope,  Linton,  Ohio. 

Thyrsidium  jasciculare  Cope,  Linton,  Ohio. 
Family  Ichthycanthidce  Moodie. 

Ichthycanthus  ohiensis  Cope,  Linton,  Ohio. 

Ichthycanthus  platypus  Cope,  Linton,  Ohio. 
Family  Molgophidee  Cope,  1875. 

Molgophis  macrurus  Cope,  Linton,  Ohio. 

Molgophis  brevicostatus  Cope.  Linton,  Ohio. 

Molgophis  wheatleyi  Cope,  Linton,  Ohio. 

Erpetobrachium  mazonensis  Moodie,  Mazon  Creek. 

Pleuroptyx  clavatus  Cope,  Linton,  Ohio. 

Phlegethontia  linearis  Cope,  Linton,  Ohio. 

Phlegethontia  serpens  Cope,  Linton,  Ohio. 


48  THE   COAL    MEASURES    AMPHIBIA    OF   NORTH   AMERICA. 

Class  Amphibia — Continued. 

Subclass  Lepospondylia — Continued. 
Order  Microsauria — Continued. 

Family  Sauropleuridas  Hay,  1902. 

Sauropleura  digitata  Cope,  Linton,  Ohio. 

Sauropleura  newberryi  Cope,  Linton,  Ohio. 

Sauropleura  scutellata  Newberry,  Linton,  Ohio. 

Sauropleura  pauciradiata  Cope,  Linton,  Ohio. 

Sauropleura  longidentata  Moodie,  Linton,  Ohio. 

Sauropleura  foveata  Cope,  Linton,  Ohio. 

Sauropleura  (Anisodexis)  enchodus  Cope,  Linton,  Ohio. 

Sauropleura  sp.,  Linton,  Ohio. 

Ctenerpeton  alveolatum  Cope,  Linton,  Ohio. 

Saurerpeton  latithorax  Cope,  Linton,  Ohio. 

Leptophraclus  obsoletus  Cope,  Linton,  Ohio. 

Leptophractus  dentatus  Moodie,  Linton,  Ohio. 

Leptophraclus  lineolalus  Cope,  Linton,  Ohio. 
Order  Temnospondylia  Zittel,  1887.    Coal  Measures  and  Permian. 
Family  Cricotidas  Cope,  1884. 

Spondylerpeton  spinatum  Moodie,  Mazon  Creek,  111. 
Family  Eryoptdce  Cope,  1882. 

Eryops  sp.  indet.,  Pitcairn,  Pa. 
Family  Macrerpetidce  Moodie,  1909. 

Macrerpeton  huxleyi  (Cope),  Linton,  Ohio. 

Macrerpelon  deani  Moodie,  Linton,  Ohio. 
Family  Anthracosauridce  Cope,  1875. 

Eosaurus  acadianus  Marsh,  Nova  Scotia. 

Eobaphctes  kansensis  Moodie,  Kansas. 

Baphetes  planiceps  Owen,  Nova  Scotia. 

Baphetes  minor  Dawson,  Nova  Scotia. 

Dendrerpeton  acadianum  Owen,  Nova  Scotia. 

Dendrerpelon  oweni  Dawson,  Nova  Scotia. 

(f)Platystegos  loricatum  Dawson,  Nova  Scotia. 

Genera  and  species  of  uncertain  position : 

Amblyodon  problematicum  Dawson,  Nova  Scotia. 
Proterpeton  gurleyi  Moodie,  Illinois. 
Brachydectes  neivberryi  Cope,  Linton,  Ohio. 
Order  Stereospondylia  Zittel,  1887.    Coal  Measures  (?)  and  Triassic. 
Family  Mastodonsauridm  Huxley,  1863. 

Mastodonsaurus  sp.  indet.,  Kansas. 

The  above  list  of  species  is  interesting  in  that  it  shows  the  diversity  of  structure 
among  the  Coal  Measures  Amphibia.  There  are  at  present  88  species  known,  many 
of  them  incompletely,  divided  among  1 7  families  and  5  orders.  The  majority  of  the 
species  are  from  the  Linton  Coal  Measures,  there  being  50  species  described  or  indi- 
cated from  these  beds.  The  Mazon  Creek  shales  have  furnished  10  species;  Nova 
Scotia  18  species;  the  remainder  of  the  species  are  from  various  localities. 


CHAPTER  VIII. 

DEFINITION  OF  THE  CLASS  AMPHIBIA  L1NNE,   1758,  DEVONIAN   TO  RECENT. 

(World-wide  distribution.) 

Cold-blooded  vertebrates;  aquatic  or  partially  terrestrial  in  habit;  body  scaled  or  naked  or 
partly  covered  with  bony  or  horny  plates;  abdomen  sometimes  protected  by  closely  packed  scutes, 
scales,  or  rods ;  skull  completely  roofed  over  or  with  a  single  vacuity ;  pterygoid-palatine  arch  complete 
or  wanting ;  stapes  always  present ;  two  occipital  condyles,  sometimes  cartilaginous ;  skull  bones  pitted 
and  grooved  by  the  lateral-line  canals,  or  smooth  and  lateral-line  canals  wanting;  parasphenoid 
well  developed ;  palatine  vacuities,  large,  small,  or  absent ;  basioccipital  partly  or  entirely  cartilaginous; 
sclerotic  plates  present  or  absent ;  mouth  always  terminal ;  teeth  sharply  conical,  smooth,  or  plicated,  with 
walls  sometimes  extremely  complicated  by  the  infolding  of  the  dentine  and  enamel.  Vertebrae  proccelous, 
opisthoccelous,  amphiccelous,  amphiplatyan,  temnospondylous,  stereospondylous,  or  cartilaginous; 
notochord  often  persistent ;  column  divisible  into  cervical,  dorsal,  and  caudal  series ;  cervical  series,  so  far 
as  known,  always  short ;  dorsal  region  long  or  short ;  a  single  sacral  or  two ;  caudal  series  short,  very  long, 
or  absent.  Pectoral  girdle  composed  of  an  osseous  scapula,  cleithrum,  clavicle,  interclavicle,  and 
coracoid  with  various  relations ;  sternum  undeveloped ;  pectoral  girdle  of  membrane  bones;  in  Triassic 
forms  producing  the  effect  of  a  plastron  on  account  of  the  high  development  of  the  clavicles  and  inter- 
clavicle. Pelvis  usually  composed  of  an  osseous  ilium  and  ischium;  pubis  when  osseous  surrounded 
by  large  amounts  of  cartilage,  usually  cartilaginous,  sometimes  calcified.  Limbs  ambulatory,  nata- 
tory, or  wanting ;  limb  bones  composed  either  entirely  of  perichondrium  or  of  perichondrium  and  a 
small  amount  of  endochondrium ;  radius  and  ulna,  and  tibia  and  fibula  free  or  fused.  Digits  3  to  5, 
usually  4  for  the  hand  and  5  for  the  foot.  Terminal  phalanges  sometimes  clawed.  Carpus  and  tarsus 
osseous  or  cartilaginous,  usually  the  latter.  Ribs  never  attached  to  a  sternal  apparatus,  single  or 
double  headed  or  intermediate,  long  and  curved  or  short  and  straight.  Articulation  with  vertebral 
column  inter-  or  intra-central.  Respiration  both  branchial  and  pulmonary;  branchiae  persistent  and 
osseous  in  some  forms.  Development  by  metamorphosis  either  in  the  egg  membrane,  on  the  back  of 
the  mother,  or  in  the  water.  No  amnion  or  allantois.  Heart  with  a  single  ventricle  and  3  or  4  pairs 
of  aortic  arches;  postcava  always  present  in  the  recent  forms. 

DEFINITION  OF  SUBCLASS  EUAMPHIBIA,  MOOD1E,   1909.    COAL  MEASURES 

TO  RECENT. 
(World-wide  distribution.) 

Moodie,  Trans.  Kans.  Acad.  Sci.,  1909,  p.  243. 

Moodie,  Geol.  Mag.,  n.  s.,  Dec.  v,  vol.  VI,  p.  220,  May,  1909. 

The  present  group  was  established  for  the  reception  of  the  Branchiosauria  and 
their  descendants,  the  Caudata,  with  the  related  forms,  the  Apoda.  The  Salientia 
are  included  provisionally,  since  there  is  no  evidence  of  the  origin  or  relationship 
of  this  group  of  animals  to  other  Euamphibia  save  that  they  have  attained  the 
same  stage  of  evolution.  They  are  in  no  way  closely  related  to  any  known  group 
of  Amphibia,  recent  or  extinct,  but  they  stand  on  the  same  plane  of  development 
as  the  Caudata  and  present  similar  structures,  i.e.,  a  single  ventricle  in  the  heart, 
external  branchiae  in  the  young,  a  glandular  skin,  perichondral  bone,  and  a  large 
parasphenoid.  The  origin  of  the  Salientia  is  a  puzzle  and  must  remain  so  until 
further  paleontological  evidence  is  forthcoming.  Wyman,  Cope,  and  the  writer 
have  all  remarked  on  the  similarity  of  structure  between  the  Salientia  and  the 
single  known  specimen  of  Pelion  lyelli  Wyman  from  the  Linton,  Ohio,  Coal  Measures. 

49 


50  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

The  subclass  Euamphibia  may  be  defined  as  follows:  Aquatic  or  terrestrial 
Amphibia;  development  by  metamorphosis;  external  branchiae  present  in  the  young; 
bones  almost  entirely  perichondral ;  carpus  and  tarsus  never  ossified ;  osseous  pubis 
absent;  vertebrae  usually  amphiccelous  with  persistent  notochord;  ribs  short  and 
straight  or  flat  and  slightly  curved,  or  absent;  digits  4  in  the  hand  and  5  in  the 
foot;  skull  never  grooved  or  pitted,  nor  cut  by  the  lateral-line  canals;  lateral-line 
organs  present  in  the  skin ;  sclerotic  plates  present  or  absent ;  tail  long  and  flattened 
or  absent.    Ribs  in  Triton  walthi  are  secondarily  long  and  curved. 

DEFINITION  OF  THE  ORDER  BRANCHIOSAURIA  LYDEKKER,  1890.    COAL  MEASURES 
AND  PERMIAN  OF  NORTH  AMERICA  AND  EUROPE. 

Lydekker,  Cat.  Fossil  Reptilia  and  Amphibia,  pt.  iv,  p.  208,  1890. 

Extinct,  salamander-like  amphibians,  with  broad,  obtusely  rounded  cranium; 
external  branchiae  present  in  young;  sclerotic  plates  present;  bones  of  the  cranium 
not  ornamented  with  deep  pits  and  grooves  nor  cut  by  the  lateral-line  canals,  though 
sometimes  ornamented  with  slight  scorbiculations ;  notochord  always  persistent; 
vertebrae  cartilaginous  (in  caudal  region)  or  but  partially  ossified,  the  ossification 
being  entirely  perichondral ;  a  single  sacral  vertebra ;  transverse  process  of  vertebrae 
large  in  dorsal  region;  ribs  always  short,  straight,  and  heavy,  present  throughout 
the  length  of  the  vertebral  column  and  borne  on  the  transverse  processes  centrally ; 
caudal  region  of  moderate  length  with  elongate  fleshy  tail ;  usually  20  presacrals, 
of  which  4  or  5  may  be  considered  cervicals;  limbs  natatory  and  always  present, 
well  developed;  elements  of  the  appendicular  skeleton  composed  entirely  of  peri- 
chondrium; carpus  and  tarsus  cartilaginous;  digits  4  in  the  hand  and  5  in  the  foot; 
phalangeal  formula  for  the  hand  usually  2-2-3-2  and  for  the  foot  2-3-4-3-2 ;  distal 
phalanges  not  clawed;  abdomen  covered  with  closely  packed  corneous  scutes  or 
scales;  body  naked  or  covered  with  minute  horny  scales;  median  and  dorsal  lateral 
lines  present  on  the  posterior  part  of  body  and  on  tail. 


CHAPTER  IX. 

THE  AMERICAN  COAL  MEASURES  BRANCHIOSAURID/E. 

Definition  of  the  Family  Branchiosaurid^b  Fritsch,  1879.      Coal  Measures  and 

Permian  of  North  America  and  Europe. 

Fkitsch,  Fauna  der  Gaskohle,  13d.  I,  p.  69,  fig.  30,  1879. 

Lydekker,  Cat.  Fossil  Rcptilia  and  Amphibia,  pt.  iv,  p.  210,  1890  (Protritonidae). 

Stegocephalic,  salamander-like  animals,  with  broad,  anteriorly  truncate  skull. 
Teeth  smooth  with  large  pulp-cavity.  The  parasphenoid  narrowed  anteriorly,  pos- 
teriorly expanded  to  a  shield-shaped  plate.  Vertebrae  with  the  notochord  persistent 
and  intravertebrally  expanded.  Pelvis  well  developed,  the  ilium  and  ischium  osse- 
ous with  large  cartilaginous  margins,  the  pubis  unknown,  possibly  hyaline  cartilage. 
Ribs  short,  straight,  present  on  almost  all  vertebras.  Skin  with  delicately  ornamented 
scales.  Eyes  with  sclerotic  plates.  Palatal  elements  toothless  or  with  small  tooth- 
like tubercles  on  pterygoids  and  palatines.  Ventral  armature  on  throat,  chest,  and 
abdomen,  extending  on  to  the  limbs,  consisting  of  small  delicate  scutellae  arranged 
in  a  chevron  pattern. 

The  above  definition  is  modified  from  Fritsch  (Fauna  der  Gaskohle,  Bd.  I,  p.  69, 
1879). 

The  North  American  species  are:  (?)  Sparodus  sp.  indet.  Dawson,  Micrerpeton 
caudatum  Moodie,  Mazonerpeton  longicaudatum  Moodie,  Mazonerpeton  costatum 
Moodie,  Eumicrer peton  parvum  Moodie. 

Genus  MICRERPETON  Moodie. 

Moodie,  Jour.  Geol  ,  17,  p.  39,  figs.  1  to  6,  1909. 

Type  Micrerpeton  caudatum  Moodie. 

The  genus  Micrerpeton,  of  which  the  single  species  is  described  below,  was  the 
first  evidence  of  the  occurrence  of  the  Branchiosauria  in  America.  There  have  been 
three  other  genera  referred  to  the  Branchiosauria  from  North  American  deposits, 
but  there  is  good  evidence  that  none  of  them  belong  there.  The  genus  Amphibamus 
was  originally  referred  to  the  Xenorhachia  by  Cope  (105,  pp.  134-137)  on  account  of 
the  supposed  cartilaginous  condition  of  the  vertebrae  and  the  absence  of  ribs.  Later 
he  abandoned  this  order  and  placed  the  form  in  the  Branchiosauria,  where  it  was 
retained  by  Zittel  (642).  Recently  Hay  has  shown  (316),  and  I  am  able  to  corrob- 
orate his  statement,  that  ribs  are  present  in  the  species,  and  that  they  are  long  and 
curved,  not  at  all  like  the  short  ribs  of  the  true  Branchiosauria.  These  long,  curved 
ribs  undoubtedly  exclude  Amphibamus  from  the  Branchiosauria  and  indicate  its 
close  affinities  with  the  Microsauria.  The  genus  Pelion  has  also  been  referred  to  this 
order  on  purely  negative  evidence  (642,  p.  375).  This  genus  is  excluded  from  the 
Branchiosauria  by  the  well-ossified  condition  of  the  limb  bones,  in  which  the  endo- 
chondral ossification  is  seen  to  be  well  developed,  a  condition  not  found,  so  far, 
among  the  true  Branchiosauria.  The  form  of  the  head  and  the  elongate  hind  limb 
would  also  tend  to  exclude  the  genus  from  the  Branchiosauria.  In  the  Branchiosauria 

51 


52 


THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


the  fore  limb  is  usually  larger  than  the  hind  limb,  the  reverse  of  which  is  the  case  in 
Pelion.  The  genus  Sparodus,  as  it  occurs  in  North  America,  is  uncertain.  It  is 
indicated  by  remains  which  are  almost  impossible  of  determination. 

The  genus  Micrerpeton  may  be  distinguished  from  other  known  Branchiosauria  by 
the  large  size  and  anterior  position  of  the  orbits,  absence  of  a  posterior  table  to  the 
skull,  the  short,  heavy  limb  bones,  the  slender  ilium,  and  the  expanded,  elongate,  and 
laterally  compressed  tail.  The  genus  may  be  defined  as  follows :  Small  forms,  the 
known  representative  attaining  a  length  of  less  than  2  inches ;  head  broad  and  short ; 
sclerotic  plates  present ;  interorbital  space  less  than  the  least  diameter  of  the  orbit ; 
occiput  concave;  pineal  foramen  in  the  line  which  cuts  the  posterior  edge  of  the 
orbits;  teeth  small,  pleurodont  denticles;  presacral  vertebrae  20  or  21,  of  which 
probably  5  are  cervical;  1  sacral;  ribs  short,  straight,  and  heavy,  central;  scapula 
ovoid ;  limbs  stumpy  and  heavy,  fore  limb  exceeding  the  hind  in  size ;  endochondral 
ossifications  distinctly  absent;  tail  long,  expanded,  and  flattened,  probably  provided 
with  a  thin  expanded  membrane;  body  covered  with  minute,  ovoid  or  rounded 
scales  which  are  ornamented  with  concentric  lines ;  color  markings  vertical  to  the 
long  axis  of  the  body  and  abundantly  present  on  the  tail ;  lateral-line  organs  repre- 
sented by  the  dorsal  and  median  lateral  lines  on  the  tail,  the  sensory  pits  probably 
occurring  in  specialized  darkened  scales.  Coal  Measures  of  Mazon  Creek  shales 
near  Morris,  Grundy  County,  Illinois. 

Micrerpeton  caudatum  Moodie. 
Moodie,  Jour.  Geol.,  17,  p.  39,  figs.  1-6,  1909. 

Type:  Specimen  No.  12,313,  Walker  Museum,  University  of  Chicago. 

Horizon  and  locality :  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  species  is  represented  by  very  complete  remains  (plate  2),  which  are  pre- 
served on  opposite  halves  of  a  nodule.  The  specimen  was  collected  many  years  ago 
by  Mr.  W.  F.  E.  Gurley  at  Mazon  Creek,  but  it  has  never  before  been  studied, 
although  Dr.  Newberry  examined  it  and  said  in  a  note  that  Professor  Cope  should 
see  it.  Unfortunately  Cope  did  not  see  it  and  it  lay  unknown  for  more  than  a  quar- 
ter of  a  century.  I  am  indebted  to  Dr.  Stuart  Weller  for  calling  my  attention  to  the 
specimen,  as  well  as  for  the  privilege  of  describing  it. 

The  specimen  is  exceptionally  perfect  (plate  25,  fig.  4).  Nearly  all  the  skeletal 
elements  are  present,  and  the  general  contour  of  the  body,  the  character  of  the  dermal 
covering,  the  color-markings,  the  lateral-line  system,  and  many  other  features  of 
interest  have  been  detected.  Such  completeness  of  preservation  is  very  uncommon 
even  among  the  remains  obtained  from  this  locality.  In  this  case  the  entire  form 
was  preserved,  but  the  collector,  in  cracking  the  nodule,  lost  the  chips  containing 
the  feet,  so  that  only  portions  of  the  limbs  remain.  It  is  thus  impossible  to  deter- 
mine the  phalangeal  formula,  but  the  feet  were  probably  like  those  of  Branchio- 
saurus  amblystomus  Credner,  as  given  by  Credner,  to  which  species  the  present  form 
is  closely  allied  and  indeed  must  be  placed  in  the  same  family  with  Branchiosaurus, 
Pelosaurus,  and  Melanerpeton. 

The  remains  here  described  represent  a  small,  salamander-like  form,  and  they 
are  among  the  earliest  geological  evidence  of  the  group,  which,  without  doubt,  gave 


PLATE   2 


Orbits  blackened  by 
pigment 


L-..  Pineal  eye 
■  ..Mandible 


Interclavicle 


Spines  of  vertebrae 


Centra  of  vertebje 


Tibia 


Femur , -' 


Impression    of    the  fleshy    and. 

membranous    parts    of 

the  tail. 


Banded  color  markings 
on  tail 


Impressions  of  cartilagi- 
nous vertebra 


_'— .  Dorsal  lateral  line 


Median  lateral  line 


Drawing  X  3. 5.  of  type  specimen  of  Micrerpeton  caudatum  Moodie,  from 
the  Coal  Measures  of  Mazon  Creek,  Illinois,  showing  skeletal  elements, 
form  of  head  and  tail,  the  lateral-line  sense  organs,  banded  color- 
markings,  and  ventral  scutellse.  On  the  edges  of  the  tail  impression 
are  indications  that  in  life  the  tail  had  a  thin  fold  of  skin  above 
and  below  the  fleshy  portion,  much  as  in  the  larva;  of  .  I ni/dys/oitia  at 
the  present  day. 


THE   AMERICAN   COAL   MEASURES   BRANCHIOSAURID^E.  53 

rise  to  the  modern  salamanders.  The  parts  preserved  in  the  specimen  are:  the 
complete  outline  of  the  head  with  most  of  the  cranial  elements  clearly  distinguish- 
able, as  well  as  the  black  pigment  of.  the  choroid;  the  entire  vertebral  column,  in- 
cluding pits  in  the  tail  region,  where  the  vertebrae  were  without  doubt  entirely  car- 
tilaginous; parts  of  the  pectoral  girdle;  the  ilium;  the  left  humerus;  the  ventral 
scutellation;  the  ribs  of  one  side  of  the  body  and  indications  of  ribs  on  the  other; 
portions  of  both  hind  limbs;  and  a  complete  impression  of  the  fleshy  tail.  On  this 
impression  of  the  tail  are  preserved  small,  horny  scales,  transverse  color-markings, 
and  the  distinct  impressions  of  the  lateral-line  system. 

The  bones  of  Micrerpeton  caudatum,  as  in  so  many  of  the  fossils  from  this  local- 
ity, have  been  replaced  by  a  white,  friable  mineral  which  is  probably  kaolin.  The 
animal  is  preserved  on  its  back  and  it  is  thus  illustrated  from  the  ventral  side.  The 
entire  length  of  the  animal  is  only  49  mm.,  of  which  the  tail  occupies  nearly  half. 

The  head  has  much  the  same  shape  as  in  the  species  of  Branchiosaurus  described 
and  figured  by  Fritsch  (251),  Credner  (181),  and  Theyenin  (568).  The  eyes  occupy 
relatively  the  same  position  as  in  that  genus.    The  orbits  are  very  large  and  broadly 


Fig.  13. — Restoration  of  Micrerpeton  caudatum,  a  branchiosaur 
from  the  Coal  Measures  of  Illinois.    X  2. 

oval.  Within  the  borders  of  the  rim  the  stone  is  blackened  as  though  by  the  black 
pigment  of  the  iris,  such  as  Cope  has  described  in  Amphibamus.  Under  a  rather 
high  power  of  magnification  the  cranial  bones  are  seen  to  be  represented  by  mere 
flakes  of  white  mineral  matter.  The  sutures  separating  the  cranial  elements  are 
distinctly  preserved  on  the  main  half  of  the  nodule. 

The  openings  of  the  skull  are  five — the  two  orbits,  the  two  minute  nostrils,  and 
the  pineal  foramen.  A  median  suture  separates  the  skull  into  halves  and  the  pineal 
foramen  lies  slightly  anterior  to  the  posterior  third  of  its  length.  The  boundaries 
of  the  premaxillas  are  not  distinct,  but  they  are  very  small  elements  and  form  the 
inner  border  of  the  nostrils,  which  are  clearly  indicated  by  bosses  of  stone.  The 
nasal  element  is  nearly  square  and  lies  anterior  to  the  frontal,  which  it  borders 
broadly.  The  parietal  is  about  the  same  size  as  the  frontal  and  it  apparently  forms 
a  portion  of  the  inner  border  of  the  orbit,  although  this  is  not  assured.  The  parietal 
is  elongate  and  unites  posteriorly  with  the  postparietal.  The  postparietal,  with  the 
tabulare  and  the  squamosal,  form  the  posterior  boundary  of  the  skull,  and  they  are 
hence  not  unlike  the  same  elements  in  other  Stegocephala.  The  prefrontal  forms 
the  anterior  border  of  the  orbit.    The  lacrimal  has  not  been  detected.    The  maxilla 


54  THE  COAL  MEASURES  AMPHIBIA   OF   NORTH   AMERICA. 

is  elongate  and  forms  the  antero-lateral  border  of  the  skull.  The  jugal  forms  an 
important  element  in  the  lateral  border  of  the  cranium  and  joins  the  quadratojugal 
posteriorly.  The  postfrontal  is  triangular  and  with  the  postorbital  forms  the  pos- 
terior border  of  the  orbit.  Both  of  the  elements  are  acuminate  posteriorly,  although 
the  suture  between  them  is  indistinct,  and  they  inclose  between  their  posterior  acumi- 
nations  an  anterior  projection  of  the  supratemporal.  The  squamosal  has  the  usual 
relations  and  borders  the  supratemporal  laterally.  The  latter  element  forms  the 
quadrate  angle  of  the  cranium. 

The  entire  length  of  the  vertebral  column  is  preserved,  although  the  nature  and 
structure  of  its  elements  can  not  be  determined.  The  impressions  of  a  few  of  the 
vertebrae  show  that  some  of  the  centra  were  amphiccelous,  but  other  than  this 
nothing  is  definite.  The  cavities  which  the  centra  occupied  were  filled  by  the  white 
mineral  matter  and  the  force  of  the  blow  which  cracked  the  nodule  destroyed  the 
form  of  the  mold.  It  is  possible  that  where  the  mineral  matter  has  filled  the  cavities 
the  centra  were  osseous  or  partly  cartilaginous,  and  where  the  cavities  were  unfilled 
the  centra  were  entirely  cartilaginous.  The  length  of  the  vertebral  column  from  the 
base  of  the  skull  to  the  last  impression  of  a  cartilaginous  centrum  is  33  mm. 

The  number  of  centra  between  the  sacral  vertebra  and  the  skull  is  20  as  they 
are  preserved,  but  there  may  have  been  one  more,  the  atlas.  Fritsch  has  represented 
21  in  his  restoration  of  Branchiosaurus  salamandroides,  and  this  is  a  further  indica- 
tion of  an  affinity  between  the  two  genera,  although  Credner  has  represented  26  pre- 
sacral vertebras  in  Branchiosaurus  ambly stomas.  The  presacral  vertebrae  are  thus 
seen  to  vary  within  narrow  limits,  but  the  number  of  presacrals  is  near  20,  and  this 
may  be  taken  as  typical.  It  is  interesting  to  notice  that  in  modern  forms  of  the 
salamanders  the  presacral  vertebrae  number  about  20.  There  is  but  a  single  sacral 
centrum  in  Micrerpeton.  The  sacral  rib  has  not  been  detected,  but  it  is  restored 
after  the  condition  found  in  Branchiosaurus.  The  right  femur  partially  covers  the 
sacral  vertebra,  and  its  structure  can  not  be  determined.  I  count  impressions  of  17 
caudal  centra,  of  which  at  least  12  may  have  been  partially  ossified.  In  the  cervi- 
cal region  there  are  distinct  impressions  of  transverse  processes  on  at  least  5  verte- 
brae, and  this  number  is  assigned  to  the  neck,  although  it  is  by  no  means  certain 
that  this  is  the  correct  number.  The  neck  was  at  least  short,  if  we  may  judge  from 
the  position  of  the  remains  of  the  pectoral  girdle.  No  cervical  ribs  are  definitely 
determined.  There  is  a  short  rib  lying  between  the  fifth  and  sixth  vertebrae,  but  to 
which  it  belongs  is  uncertain. 

There  are  impressions  of  10  ribs  preserved  on  one  side  of  the  vertebral  column 
and  one  on  the  other  side.  They  are  short,  straight,  and  heavy,  as  are  the  same  ele- 
ments in  Branchiosaurus.  This  character  alone  is  sufficient  to  place  Micrerpeton 
among  the  Branchiosauria,  since  no  such  ribs  are  known  in  other  groups  of  the 
extinct  amphibians.  The  ribs  preserved  lie  next  the  seventh  to  the  seventeenth 
vertebrae  on  the  left  side,  and  there  is  one  on  the  right  side  which  may  belong  to  either 
the  fifth  or  sixth  vertebra.  They  are  central  in  their  attachment,  and  in  this  they 
agree  well  with  the  mode  of  rib  attachment  in  the  modern  salamanders.  All  of  the 
ribs  are  single-headed  and  are  composed,  for  the  most  part,  of  perichondral  tissue. 


THE   AMERICAN   COAL   MEASURES   BRANCHIOSAURID/E.  55 

The  position  of  the  ribs  in  the  matrix  inclined  backwards,  and,  making  a  small  angle 
with  the  vertebral  column,  is  very  suggestive  of  the  condition  in  Branchiosaurus. 

The  pectoral  girdle  is  represented  by  three  distinct  elements  of  the  left  side, 
which  are  identified  as  scapula,  clavicle,  and  coracoid,  following  the  nomenclature 
given  by  Woodward  (631),  although  Credner  (186)  would  name  them  otherwise. 
The  scapula  is  represented  by  an  ovoid  fragment  lying  next  to  the  vertebral  column. 
The  clavicle  was  probably  spatulate,  as  in  Melanerpeton,  but  the  inner  end  of  the 
element  is  not  visible.  The  coracoid  is  represented  by  its  outer  end  only,  and  its 
inner  pointed  extremity  is  not  visible.    The  interclavicle  has  not  been  detected. 

The  humerus  lies  somewhat  to  one  side  of  the  pectoral  girdle,  as  if  there  had  been 
a  large  amount  of  epiphyseal  cartilage.  Its  position  may  be  due  to  post-mortem 
shifting,  but  there  is  little  other  evidence  of  any  movement  after  deposition.  The 
humerus  is  a  large,  heavy  bone  in  comparison  with  the  rest  of  the  skeleton.  It  is 
expanded  at  each  end,  and  its  ends  show  concavities,  proving  that  the  bone  is  formed 
principally  of  perichondral  tissue,  as  would  be  expected  from  such  an  early  Branch- 
iosaurian.  The  endochondrium  has  not  yet  developed  in  this  form,  which  is  evi- 
dently adult.    There  is  no  other  element  of  the  arm  present. 

There  is  but  a  single  element  of  the  pelvis  preserved,  a  slender  elongate  rod 
which  is  undoubtedly  the  ilium,  since  it  has  the  usual  position  for  that  element  and 
is  much  too  large  for  a  sacral  rib.  It  has  much  the  same  shape  as  the  ilium  in  the 
modern  Salamandra,  and  is  not  expanded  as  is  the  ilium  of  Branchiosaurus.  This 
element,  like  the  humerus,  seems  to  have  been  but  a  hollow  cylinder  of  bone  and 
undoubtedly  had  cartilaginous  ends,  as  in  the  ilium  of  the  recent  Salamandra.  The 
two  femora  are  preserved  nearly  entire,  the  right  one  lying  upon  and  partly  obscuring 
the  sacral  vertebra.  The  femur  is  much  more  slender  than  is  the  humerus,  with 
slightly  expanded  ends,  and,  like  the  humerus,  shows  the  concavities  at  the  ends, 
indicative  of  the  perichondral  character  of  the  tissue  composing  it.  There  are 
two  elements  of  the  leg  preserved  more  or  less  entire.  The  larger  bone  may  repre- 
sent the  tibia  and  the  smaller  the  fibula.  They  both  present  characters  similar  to 
those  of  the  femur  and  humerus.  They  are  simple  rods  of  bone  tapering  at  the  distal 
end.    The  feet  have  been  lost,  though  doubtless  they  were  present  at  one  time. 

The  ventral  surface  of  the  body,  as  in  other  members  of  the  Branchiosauria,  was 
covered  and  protected  by  a  series  of  small  scutes  arranged  in  the  regular  chevron 
pattern.  The  form  of  the  scutes  and  their  number  can  not  be  determined.  The  lines 
which  represent  them  are,  however,  distinct.  Some  of  the  scutes  are  missing  and 
some  of  them  are  obscured  by  lying  over  the  vertebral  column.  They  are  all 
somewhat  shifted  to  the  left.  The  lines  are  very  small  and  close  together.  I  count  16 
of  them  in  a  distance  of  2  mm.  In  length  the  longest  line  preserved  is  a  little  more 
than  4  mm.,  measuring  from  the  point  of  the  chevron.  The  lines  representing  the 
scutes  come  to  a  point  in  a  median  ridge  which  is  now  represented  by  a  line.  The 
dermal  scutes  on  the  abdomen  were  probably  the  forerunners  of  the  abdominal  ribs 
of  the  reptiles  (fig.  9). 

The  impression  of  the  tail  contains  some  of  the  most  interesting  features  in  the 
entire  specimen.    Scattered  over  it  and  in  places  laid  in  mosaic  are  impressions  of 


56  THE    COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

small  dermal  scales,  which  may  have  covered  the  entire  body.  In  form  the  scales 
are  ovoid,  being  half  as  wide  as  long,  and  the  markings  on  the  scales  partake  of  the 
nature  of  radiating  lines,  much  after  the  pattern  of  the  sculpturing  of  the  cranial 
bones  in  the  Microsauria.  The  scales  are  less  than  0.5  mm.  in  diameter  and  their 
character  can  only  be  ascertained  under  high  magnification.  Near  the  middle  of 
the  tail  there  are  preserved  distinct  transverse  bands  of  dark  color,  which  are  more  or 
less  evident  throughout  the  entire  tail  impression,  but  they  are  elsewhere  not  so 
distinct  as  in  the  central  region.  The  lines  are  evidently  due  to  rows  of  pigmented 
scales,  and  in  all  probability  the  animal's  body  was  transversely  striped. 

The  most  interesting  and  important  single  structure  discovered  on  the  specimen 
is  the  impression  of  the  lateral-line  system,  which  is  clearly  evident  as  two  dark 
lines  on  the  impression  of  the  fleshy  part  of  the  tail.  The  sense-organs  are  repre- 
sented by  two  longitudinal  rows  of  pigmented  scales,  one  beginning  at  the  tip  of  the 
tail,  the  other  taking  its  origin  from  the  median  line  somewhat  further  forward.  I 
am  indebted  to  Dr.  Katashi  Takahashi  for  calling  my  attention  to  the  similarity  of 
this  arrangement  to  that  found  in  the  recent  Necturus.  The  arrangement  and  dispo- 
sition of  the  lines  containing  the  sense-organs  is  practically  the  same  in  the  two 
forms.  The  median  lateral  line  takes  its  origin  from  the  extreme  tip  of  the  tail  and 
is  continued  to  the  base,  where  the  impression  is  broken.  The  dorsal  lateral  line 
has  its  origin  rather  abruptly  from  the  median  lateral  line  at  a  distance  of  6  mm.  from 
the  tip  of  the  tail.  The  sense-organs  were  undoubtedly  located  beneath  specialized 
pigmented  scales  on  the  surface,  and  to  this  pigment  is  due  the  preservation  of 
the  lines. 

The  fact  that  the  arrangement  of  the  sense-organs  of  Micrerpeton  corresponds  so 
exactly  to  the  condition  found  in  Necturus  is  of  considerable  interest.  Necturus 
alone  among  the  modern  tailed  Amphibia  has  the  arrangement  described  for  the 
lateral-line  system  of  Micrerpeton.  All  other  forms  of  the  Caudata,  as  also  the  larval 
forms  of  the  Salientia,  have  an  arrangement  of  the  lateral-line  system  which  is  per- 
fectly distinct  from  that  found  in  Necturus,  although  the  basis  of  the  same  arrange- 
ment is  found  in  all.  In  Ambly stoma,  for  instance,  the  median  lateral  line  is  not 
present  on  the  tail,  and  the  dorsal  line  is  incompletely  developed.  The  close  simi- 
larity of  the  arrangement  of  the  systems  of  sense-organs  in  the  two  forms,  Micrer- 
peton and  Necturus,  may  be  of  genetic  significance  with  regard  to  the  latter  form. 
The  lateral-line  sense-organs  are  of  a  very  fundamental  significance,  and  it  is  not  at 
all  improbable  that  the  same  arrangement  of  the  lines  has  existed  from  the  Carbo- 
niferous period  or  earlier.  We  know  that  such  has  been  the  case  in  a  great  many  of 
the  fishes.  The  ancestors  of  the  modern  Caudata  must  have  originated  somewhere 
in  the  basal  Carboniferous  or  earlier  periods,  and,  in  the  writer's  opinion,  the  Branch- 
iosauria  represent  the  ancestral  group  of  the  Caudata.  This  suggestion  is  by  no 
means  new,  since  Baur  and  others  have  held  the  same  view.  This  topic  has  been 
discussed  at  length  elsewhere  (459)  by  the  writer. 

The  relations  of  the  form  Micrerpeton  caitdatum  are  readily  determined.  The 
number  of  the  presacral  vertebrae,  the  form  and  position  of  the  ribs,  the  shape  of  the 
skull,  the  arrangement  of  the  cranial  elements,  the  structure  of  the  pectoral  girdle 


THE   AMERICAN   COAL   MEASURES   BRANCHIOSAURID/E.  57 

and  the  character  of  the  ventral  armature  all  clearly  bespeak  a  close  relationship 
with  Branchiosaurus,  Melanerpeton,  Pelosaurus,  and  other  European  branchiosau- 
rian  forms  from  the  Upper  Carboniferous  and  Lower  Permian. 

The  above-described  species,  with  others  given  below,  is  the  earliest  geological 
evidence  of  the  Branchiosauria,  since  the  oldest  European  forms  are  from  the  Ste- 
phanian  (Upper  Carboniferous),  which  probably  lies  somewhat  above  the  horizon  of 
the  Allegheny  series  of  North  America.  The  presence  of  the  Branchiosauria  in 
America  is  of  considerable  interest  in  the  bearing  it  has  on  the  distribution  and 
migration  of  the  Paleozoic  animals.  Knowledge  of  how  the  group  came  to  occur 
in  such  widely  separated  localities  in  approximately  contemporary  geological  strata  is 
an  unsolved  problem  of  paleontology.  It  is  possible  that  the  piscian  ancestors  of  the 
Amphibia  migrated  across  or  along  the  borders  of  the  seas  and  began  the  amphibian 
phase  of  development  independently  in  the  two  continents.  That  evolution  should, 
in  this  case,  have  followed  almost  exactly  parallel  lines  seems  incredible. 


Measurements. 
ram. 


Length  of  entire  animal 49  Length  of  rib  1.5 

length  of  head  in  median  line 6.5  Length  of  scapula  3 

Width  of  head  at  posterior  border   8-  Maximum  width  of  clavicle   2 

Length  of  orbit  2.5  Length  of  humerus   2.5 

Width  of  orbit   2  Length  of  ilium  1.5 

Interorbital  space 2  Length  of  femur  2 

Length  of  the  vertebral  column   33  Length  of  tibia   ....  1.5 

Length  of  the  vertebral  centrum  in  dorsal  series.  .  0.5  Length  of  tail  impression  21.5 

Length  of  trunk  from  base  of  skull  to  sacrum.  ...  22  Width  of  tail  impression  at  base   4 

Genus  EUMICRERPETON  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  330,  1912. 

Type:  Eumicrerpeton  parvum  Moodie. 

The  genus  is  established  on  three  well-preserved  specimens  representing  nearly 
the  entire  anatomy.  The  generic  characters  are  found  in  the  very  broad  posterior 
table  of  the  skull,  with  its  short  length,  reduction  of  tympanic  notch,  and  shortness 
of  body.  The  body-length  of  Eumicrerpeton  (plate  5,  fig.  1)  is  less  proportion- 
ately than  that  of  other  closely  allied  genera.  Other  generic  characters  are  found  in 
the  sharp  postero-lateral  angle  of  the  skull,  and  it  is  to  be  distinguished  from  Micrer- 
peton,  especially,  by  the  short,  stumpy  limb  bones.  The  narrow,  elongate  eye, 
placed  close  to  the  edge  of  the  skull,  is  a  character  not  observed  hitherto  in  the 
Branchiosauria.    The  genus  is  closely  allied  to  Branchiosaurus  of  Europe. 

Eumicrerpeton  parvum  Moodie. 

Moodie,  Proc.  U.  S.  Nat.  Mus.,  40,  p.  430,  fig.  1,  1911. 

Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  pp.  331-336,  p'-  3.  figs-  3  and  4;  pi.  4;  pi.  5,  fig.  1;  pi.  6,  figs.  1 

and  2,  1912. 
Moodie,  Amer.  Nat.,  44,  pp.  367-375,  figs.  1-4,  1910. 
Moodie,  Science,  n.  s.,  xxxi,  No.  789,  p.  233. 

Type:  Specimen  No.  803,  Yale  University  Museum.  Other  specimens,  No. 
802,  Yale  University  Museum,  and  No.  4400,  U.  S.  National  Museum. 

Horizon  and  locality:  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  impression  of  the  outline  of  the  entire  body  is  preserved  (plate  3,  figs.  1  and  2) 
in  three  specimens,  and  in  all  are  found  molds  and  impressions  of  the  alimentary 


58  THE   COAL   MEASURES   AMPHIBIA    OF   NORTH    AMERICA. 

canal,  which,  in  one  specimen  (471),  are  remarkably  complete  and  instructive.    The 
three  specimens  will  be  discussed  separately,  since  they  show  different  features. 

The  impression  of  the  larger  animal  (No.  803,  Yale  University  Museum),  which 
is  probably  an  adult,  presents  the  following  elements:  the  entire  skull,  both  humeri, 
impressions  of  posterior  and  anterior  ventral  armature,  portions  of  the  alimentary 
canal,  one  femur,  portions  of  a  fibula  and  tibia,  and  the  entire  impression  of  the  tail, 
on  which,  as  in  Micrerpeton  caudatum,  there  occur  two  definite  dark  lines,  one  begin- 
ning at  the  tip  of  the  tail  and  running  obliquely  along  the  tail  to  where  the  impres- 
sion is  broken  at  the  anal  region;  the  other  beginning  at  a  distance  of  4.5  mm.  from 
the  tip  and  running  almost  parallel  with  the  median  line.  These  two  lines  undoubt- 
edly represent  the  lateral -line  system. 

The  skull  is  especially  noted  for  its  shortness  and  the  great  posterior  width,  as 
well  as  for  the  almost  entire  absence  of  the  tympanic  notch.  The  pineal  foramen  is 
located  on  a  line  with  the  posterior  border  of  the  orbits.  The  eyes  themselves  are 
narrow  and  acuminate  at  each  end,  with  a  pronounced  convexity  inwards  and  a 
flattening  on  the  outer  margin.  They  are  located  on  the  very  border  of  the  skull, 
but  relatively  more  posterior  than  in  Micrerpeton.  No  sclerotic  plates  are  evident. 
The  median  suture  can  be  indistinctly  observed  running  the  entire  length  of  the 
skull.  The  sutures  bounding  the  outside  of  the  frontals  and  the  squamosals  are 
partially  evident,  but  not  satisfactorily  preserved.  The  mandible  is  represented  by 
a  mold  which  in  wax  impression  shows  short,  stumpy  teeth. 

Posterior  to  the  skull  at  a  distance  of  a  millimeter  there  are  two  sharp  impres- 
sions which  may  represent  the  anterior  edges  of  the  interclavicle  or  they  may  be 
branchial  elements.  They  are  distinctly  curved,  however,  and  probably  represent 
portions  of  the  interclavicle.  A  wax  impression  does  not  show  a  discrete  structure, 
but  the  boundaries  of  some  larger  element.  No  other  remains  of  the  pectoral  girdle 
can  be  discerned.  The  humeri  are  short  and  relatively  thick.  Wax  impressions 
show  them  to  have  had  truncate  or  slightly  concave  ends,  thus  indicating  the 
absence  or  slight  development  of  the  endochondrium.  No  other  elements  of  the  arm 
are  preserved. 

The  impression  of  an  elongate  femur  and  the  heads  of  the  tibia  and  fibula  of  the 
left  side  are  preserved. 

The  ventral  armature  is  preserved  in  two  small  patches,  and  these  show  the 
chevron-shaped  rods  to  have  been  very  fine — much  more  delicate  than  in  Micrerpeton. 

The  body  impression  is  very  instructive  and  interesting,  both  in  showing  the 
form  of  the  body  and  because  in  it  are  preserved  the  larger  portions  of  the  alimen- 
tary canal.  The  form  of  the  body  can  best  be  discerned  by  reference  to  the  figures 
(plate  3,  figs.  1  and  2;  plate  5,  fig.  1). 

The  portions  of  the  alimentary  canal  preserved  consist  of  the  greater  portion  of 
the  stomach,  three  coils  or  loops  of  the  small  intestine,  the  rectum,  and  a  pit  which 
undoubtedly  represents  the  anal  opening.  The  anus  is  found  at  a  distance  of  1 6  mm. 
from  the  tip  of  the  tail  and  is  somewhat  removed  from  the  body  portion,  as  in  mod- 
ern salamanders.  On  each  side  of  the  posterior  end  of  the  rectum  there  occur  a  pair  of 
enlargements  which  probably  represent  the  oviducts  at  their  posterior  ends  (fig.  15,  C) . 


MOODIE 


PLATE  I 


1.  The  larger  specimen  of  Eumicrerpeton  parvum  Moodie.      XI. 

2.  The  smaller  specimen  of  Eumicrerpeton  parvum  Moodie.     X  ]. 

3.  Type  specimen  of  Erpetobrachium  mazonensis  Moodie.     X  1. 

4.  Type  specimen  of  Erierpeton  branchialis  Moodie.      X  1. 

5  and  6.     Type  specimen  of  Mazonerpelon  longicauJatum  Moodie.     X  1. 
7.     A  copy  of  Cope's  drawing  of  the  type  specimen  of  Amphibamus  grandiceps  Cope. 
The  original  was  destroyed  by  fire.     X  3. 


THE   AMERICAN   COAL   MEASURES    BRANCHIOSAURID^E. 


59 


€^ 


ic 


* 


The  tail  impression  is  more  acuminate  than  in  Micrerpeton,  but  shows  the  same 
structures  as  in  that  form,  i.e.,  the  lateral  lines  which  have  already  been  mentioned. 
Micrerpeton  was  a  more  rapid  swimmer  than  the  present  form  on  account  of  the 
greater  development  of  the  tail. 

The  second  specimen  of  the  species  (No.  802,  Yale  Museum)  shows  much  the 
same  character  as  the  specimen  already  described,  except  that  there  are  impressions 
of  small,  blunt  teeth  on  the  mandible.  The  two  humeri  and  the  femur  of  the  left 
side  are  preserved  and  the  interclavicle  is  represented  by  an  identical  impression,  as 
in  the  first-described  specimen.  The  tail  impres- 
sion, though  similar  in  form,  does  not  exhibit  so 
much  of  the  structure  of  the  lateral  lines  (fig.  15,  B). 

The  matter  of  especial  interest  in  connection 
with  this  second  specimen  is  the  remarkably  perfect 
preservation  of  the  alimentary  canal,  which  is  entire, 
except  for  the  very  anterior  end  of  the  oesophagus. 
The  posterior  portion  of  the  oesophagus,  which 
measures  3.5  mm. ,  is  clearly  preserved.  Its  anterior 
end  is  thrown  around  posteriorly  and  indicates  that 
this  end  was  loosened  after  death  and  became  dis- 
placed before  fossilization.  The  length  preserved 
may  represent  the  entire  oesophagus.  The  oesoph- 
agus is  constricted  before  it  enters  the  stomach, 
which  shows  the  usual  curvature  found  in  modern 
salamanders.  The  stomach  measures  6  mm.  in 
length  by  2  mm.  in  breadth,  and  consists  of  a 
single  enlargement  as  in  the  modern  Ambly stoma 
punctatum.  It  increases  in  size  somewhat  toward 
the  pyloric  end  and  then  very  gradually  constricts 
to  the  pylorus.  Three  divisions  of  the  small  intes- 
tine can  be  seen.  The  most  anterior  one,  corre-  a.  ThMmedmenatEimkrcrtttonpamm 
sponding  to  the  duodenum,  is  segmented,  as  though  S^hi^f  xY^OrigSi 
the  intestine  had  been  filled  with  food  before  inter-        in  United  states  National  Museum. 

a,  anus;  /,  femur;  ft,  humerus;  ic,  lntrr- 

ment.    The  remainder  of  the  intestine,  correspond-        clavicle ;"»«,  intestine;  m,  mandible;  or, 

,.       .,  .    .  ,  .       ,       .  .  ,  „  orbit;  st,  stomach;  (,  tibia  and  fibula. 

ing  to  the  ileum,  is  looped  in  the  iorm  01  two  figures  b.  Type  specimen  of  Amphibamus  tiwra- 
8  which  are  superimposed,  with  the  upper  portions  £K».  SJTffSL^S 
of  the  8  at  right  angles  to  each  other.  The  rec-  see  p-  *32-) 
turn  is  clearly  discernible,  though  its  lower  portion  is  somewhat  obscured  by  having 
the  lower  part  of  the  upper  loop  of  the  intestine  lying  over  it.  The  anus  lies  at  a 
distance  of  1 .5  mm.  posterior  to  the  transverse  line  from  the  upper  end  of  the  femur, 
and  is  quite  well  back  on  the  tail,  as  in  modern  salamanders.  In  this  specimen  also 
occur  two  oval  bodies  which  may  be  identified  as  the  lower  ends  of  the  oviducts, 
thus  indicating,  in  all  probability,  that  the  animal  was  a  female. 

A  dissection  of  several  species  of  modern  caudates  has  resulted  in  the  discovery 
that  the  adult  condition  of  the  alimentary  canal  of  all  the  species  dissected  (.1  mhly- 


B 


60  THE    COAL    MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

stoma  pimctatum,  Necturus  maculosus,  Diemyctylus  torosus,  D.  viridescens,  etc.)  is 
much  more  complex  than  that  exhibited  by  the  specimen  under  discussion.  A  very 
near  approach  to  the  condition  found  in  Eumicrerpeton  parvum  is  found  in  an  imma- 
ture branchiate  individual  of  Diemyctylus  torosus,  56  mm.  in  length,  from  a  fresh- 
water pond  on  Mount  Constitution,  on  Orcas  Island,  Puget  Sound,  Washington. 

The  similarity  of  the  intestinal  structure  is  of  considerable  importance  to  our" 
understanding  of  the  relationship  existing  between  the  Carboniferous  Branchio- 
sauria  and  the  modern  Caudata,  and  only  confirms  other  arguments,  offered  in 
another  place  (459),  concerning  their  immediate  relationship. 

The  branchiosaurian  affinities  of  the  present  species  are  almost  too  evident  to 
need  discussion.  The  entire  structure  is  essentially  similar  to  that  of  other  genera 
of  the  order. 

The  third  specimen  of  this  species  (No.  4400  of  the  U.  S.  National  Museum)  is 
almost  as  perfectly  preserved  as  were  the  other  two  specimens.  The  skull  structure, 
the  intermediate  position  of  the  pineal  foramen,  the  epiotic  notch,  and  the  shape  of 
the  skull  are  essentially  similar  to  the  described  specimens  of  the  species.  The 
present  specimen  is  more  developed  than  the  other  two  and  probably  represents 
an  adult.  The  alimentary  canal  is  perfectly  preserved. 

It  is  nearly  half  again  as  long  as  the  smallest  of  the  above-described  specimens,  and 
the  skull  is  proportionately  longer  and  wider.  There  is  preserved  also  an  impression 
of  the  anterior  ends  of  both  clavicles.  The  right  humerus  is  imperfectly  preserved,  as 
is  also  the  right  femur  and  tibia;  other  than  these  the  fossil  is  merely  an  impression. 

The  skull  is  so  similar  to  those  described  above  that  additional  description  is 
unnecessary.  The  pineal  foramen  is  quite  large  and  lies  on  a  line  which  cuts  the 
orbits  into  equal  longitudinal  parts.  The  interorbital  space  is  about  equal  to  the 
long  diameter  of  the  orbit.  Traces  of  sclerotic  plates  are  observed  in  the  left  orbit, 
but  they  are  quite  imperfect. 

The  alimentary  canal  (fig.  14a)  is  unlike  the  previously  described  structures,  in 
that  the  intestine  is  longer  and  more  convoluted.  It  lies  in  five  longitudinal  folds  and 
ends  in  an  enlarged  cloaca,  near  which  there  are  impressions  of  two  glands,  or  the  pos- 
terior ends  of  the  oviducts,  as  was  suggested  for  the  Yale  specimens.  The  creatures 
undoubtedly  fed  on  small  plants  and  animals  much  as  do  the  recent  salamanders. 

Measurements  of  Eumicrerpeton  parvum  Moodie. 

No.  803  (222),  Yale  University  Museum:                     mm.  No.  4400,  U.  S.  National  Museum:  mm. 

Length  of  animal 37.5  Length  of  entire  animal 45 

Length  of  skull 4.5  Length  of  skull 6 

Posterior  width  of  skull  at  table 6  Width  of  skull 9 

Long  diameter  of  eye 1.75  Transverse  diameter  of  orbit 1 .50 

Transverse  diameter  of  eye 65  Long  diameter  of  orbit 2.25 

Length  of  left  humerus 1.50  Interorbital  space 2.50 

Number  of  ventral  armature  rods  in  1  mm  ....  10  Diameter  of  pineal  foramen 50 

Length  of  femur 1.75  Length  of  body  from  back  of  skull  to  pelvis  ...   22 

Width  across  base  of  tail  impression 3.5  Oreatest  width  of  body 9 

Length  of  tail  from  base  to  tip 17  Length  of  tail  16 


No.  802  (471)  Yale  University  Museum: 


Width  of  tail  at  base 5 


Length  of  animal 30  Png&°r  humerus 3 

Length  of  skull ^4  'M8*  °f  *"»"*£  V  v: 2'5° 

Posterior  width  of  skull 5  Length  of  tibia  (fibula?) 1.75 

Length  of  oesophagus 3.5  Length  of  stomach 7 

Length  of  stomach .     6  Width  of  stomach 3 


Width  of  stomach 1 .33 

Estimated  length  of  intestine 18 

Width  across  base  of  tail  impression 2.5 

Length  of  tail  from  base  to  tip 7 


Length  of  intestine  (estimated) 5.6 

Width  of  intestine 1 


1  and  2.     Vertebrae  of  Spoil  dylerpeton  spinalum  Moodie.     X  1. 

3.  Tvpe  specimen  of  Mazonerpelon  rostatum  Moodie.     X  1. 

4.  Type  skeleton  of  Cephalerpeton  venliiarmnliim  Moodie.     X  0.9. 

5  and  6.     The  halves  of   the   nodule    containing   a   practically    complete 
skeleton  of  Ampliibamus  gramtkeps  Cope.     X  1. 
Originals  of  Above  figures  in  the  Yale  University  Museum. 


THE  AMERICAN  COAL  MEASURES   BRANCHIOSAURIOE.  6l 

Genus  MAZONERPETON  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vol.  vi,  No.  2,  p.  336,  1912. 

Type:  M.  longicaudat&m  Moodie. 

The  genus  is  distinguished  from  other  known  branchiosaurian  genera  by  the 
great  length  of  the  dorsal  region,  the  elongate  tail  (plate  5,  fig.  2),  with  its  well- 
developed  caudal  ribs,  the  reduction  of  the  tympanic  notch,  the  broad  nature  of  the 
scapula,  the  elongate  interclavicle,  and  the  slender  ilium.  The  number  of  dorsal 
vertebras  is  identical  with  that  of  Branchiosanrus  of  Saxony. 

Mazonerpeton  longicaudatum  Moodie. 
Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  337,  pi.  3,  figs.  i-2;pl.  7,fig.3;pl.  10.     1912.' 

Type:  Specimen  No.  795  (1234),  Yale  University  Museum. 

Horizon  and  locality:  Mazon  Creek  shales,  near  Morris,  Illinois.  (Plate  3,  figs. 
5  and  6.) 

The  remains  consist  of  the  following  elements:  an  incomplete  skull;  nearly  the 
entire  vertebral  column,  consisting  of  cervical,  dorsal,  sacral,  and  caudal  vertebrae, 
36  in  number;  several  ribs  preserved  on  each  side  of  the  vertebral  column;  a  portion 
of  the  ventral  armature;  the  scapulae;  a  clavicle;  the  interclavicle;  both  humeri;  the 
radius  and  ulna  of  one  side  and  the  ulna  of  the  other;  portions  of  both  hands;  the 
ilium  of  the  right  side;  both  femora,  and  a  partial  impression  of  the  left  tibia. 

The  skull  is,  unfortunately,  very  poorly  preserved.  Enough  remains,  however, 
to  determine  the  essential  characters.  The  skull  bones,  unlike  any  other  American 
branchiosaurian,  have  an  ornamentation  consisting  of  sharp  pits  and  elevations 
which  in  places  have  a  quincuncial  arrangement  and  in  others  take  the  form  of  defi- 
nite lines  of  pits  or  tubercles  similar  to  the  condition  found  in  many  of  the  Micro- 
sauria.  The  orbits  are  large  and  are  situated  back  of  the  median  transverse  line  of 
the  skull.  They  are  almost  circular  in  form  and  contain  6  elongated  sclerotic  plates 
very  closely  arranged  around  the  borders  of  the  right  orbit.  The.  plates  are  twice 
as  long  as  wide.  The  interorbital  width  is  1 .25  times  the  transverse  diameter  of  the 
orbit. 

Not  many  of  the  sutures  of  the  skull  are  discernible.  Portions  of  the  frontals, 
the  nasals,  the  prefrontals,  the  parietals,  and  the  supratemporals  can  be  identified. 
Their  arrangement  is  shown  in  figure  14a.  There  is  a  decided  posterior  table  to  the 
skull,  with  truncate  posterior  border.  The  tympanic  notch  is  shallow,  with  its 
outer  border  not  so  well  protected  as  in  Branchiosaurus. 

The  cervical  vertebrae  arc  incomplete,  but  their  number  was  4  or  5,  as  in  M i<  - 
rerpetott.  The  structure  of  the  dorsal  vertebrae  is  also  uncertain,  although  the  shape 
can  be  discerned.  The  vertebrae  are  short  and  thick,  very  unlike  the  long,  cylin- 
drical vertebrae  of  Cephalerpeton.  The  heavy  transverse  process  is  quite  evident  on 
the  best  preserved  vertebrae.  This  process  recalls  that  described  by  Credner  for  the 
Saxony  Branchiosauria.  Several  of  the  vertebrae  show  the  articulation  of  the  ribs 
with  this  process.  The  ribs  of  the  caudal  region  recall  very  strongly  those  of  Branch- 
iosaurus. They  are  quite  heavy  in  the  anterior  caudal  region  and  then  diminish 
rather  rapidly  to  the  point  where  the  tail  is  broken  and  lost. 


62 


THE   COAL   MEASURES   AMPHIBIA    OF    NORTH   AMERICA. 


They  are  quite  different  from 


The  ventral  armature  is  represented  by  a  patch  of  chevron  rods  21  mm.  in 
length.  The  rods  take  a  very  peculiar  form,  being  short  crescentic  bundles  of 
fine  rods,  hair-like  in  appearance.  In  one  of  these  bundles  I  count  5  smaller  rods. 
The  bundles  are  arranged  in  rows  similar  to  the  pattern  so  characteristic  of  the  Car- 
boniferous Amphibia,  as  described  elsewhere.  The  patch  of  ventral  armature  pre- 
served belongs  to  the  abdominal  region.  A  single  row  of  the  crescentic  bundles 
measures  11  mm. 

Both  scapulas  are  preserved  in  their  entire  form 
those  of  any  other  genus,  being  broadly 
crescentic  with  a  posterior  concavity  and 
an  anterior  protuberance.  The  anterior 
surface  of  both  scapulae  is  obscured.  Vas- 
cular foramina  occur  near  the  base  of  both 
scapulas;  there  being  three  of  them  in  the 
right  scapula,  arranged  in  the  form  of  an 
isosceles  triangle .  The  morphology  of  these 
foramina  is  uncertain.  They  have  never 
been  observed  among  the  Carboniferous 
Amphibia,  and,  so  far  as  I  am  aware, 
they  are  entirely  unknown  among  higher 
vertebrates. 

The  temnospondylous  Amphibia  of  the 
Carboniferous  and  Permian  possess,  in  the 
co-ossified  scapula-coracoid,  three  fora- 
mina, very  similar  to  the  ones  in  the  pres- 
ent form,  but  they  are  confined  to  the 
coracoidal  region  and,  in  the  Branchio- 
sauria,  as  identified  by  Credner,  the  cora- 
coid  is  a  free  element,  although  I  have 
never  been  sure  of  its  identity  among 
American  forms.  Williston  has  called  these 
foramina  the  glenoid,  the  supraglenoid,  and 
the  supracoracoid  foramina  (Journal  Geol- 
ogy, xvii,  No.  7).  They  are  not,  however, 
to  be  correlated  with  the  three  foramina 
above  mentioned,  since  in  the  Temnospon- 
dylia  the  foramina  belong  with  the  coracoid  and  not  with  the  scapula.  The 
condition  of  the  Temnospondylia  occurs  in  the  bony  fish  Xiphactinus  andax 
Leidy,  and  an  analogous  condition  obtains  in  the  reptiles,  as  in  the  Mosasaurs 
and  Dinosaurs. 

Near  the  outer  edge  of  the  right  scapula  there  is  a  large  fragment  preserved, 
which,  I  think,  must  be  the  misplaced  clavicle.  It  is  obscurely  triangular  or,  more 
exactly,  spatulate.  The  interclavicle  is  represented  by  fragments  only,  and  seems  to 
have  had  a  narrow  form. 


I 


I  W 


f 


\<^  Si 


Fig. 


14a. — Skeleton  of  Mazonerpeton  longicaudatum 
Moodie.  c,  carpus;  d,  clavicle;  cr,  caudal  ribs;  cv, 
caudal  vertebrae;  h,  humerus;  /,  femur;  or,  orbit; 
r,  radius;  sf>,  sclerotic  plates;  sc,  scapula;  u,  ulna; 
us,  ventral  scutelke.  From  Mazon  Creek.  Original 
in  Yale  University  Museum. 


THE   AMERICAN   COAL    MEASURES    BRANCHIOSAURID/E. 


63 


The  humeri  recall  those  of  Micrerpeton.  They  are  somewhat  elongate  and  appar- 
ently cylindrical  in  their  normal  condition,  although  somewhat  flattened  in  the  fos- 
sil. The  shaft  is  considerably  constricted  at  the  middle  and  the  ends  are  expanded, 
in  which  expansion  the  lower  end  exceeds.  The  ends  are  abruptly  truncate,  indi- 
cating a  small  amount  of  endochondral  ossification  or  its  entire  absence. 

The  mesopodial  elements,  unlike  what  is  described  for  Cephalerpeton,  are  quite 
dissimilar  in  form,  recalling  the  condition  in  Mesosanrus  brasiliensis  McGregor.  The 
larger  element  is,  apparently,  the  ulna.  It  has  the  lower  end  greatly  expanded  and 
the  shaft  is  curved  outward,  resembling  very  much  a  reptilian  ulna.  The  radius 
is  much  smaller  than  the  ulna,  lacks  the  lower  expansion,  and  is  shorter  by  i  mm. 

The  carpus  is  represented  merely  by  a  blank  space,  with  evidences  of  impres- 
sions of  cartilage  in  the  sandstone.  The  hand  of  the  right  side  contains  4  phalanges. 
There  are  2  phalanges  preserved  in  the  first  digit,  including  a  sharp-pointed  ter- 
minal phalanx,  and  the  second  digit  has  only  the  metacarpal.  The  third  has  the 
metacarpal  and  the  first  phalanx,  which  does  not  differ  in  form,  but  only  in  size, 
from  the  metacarpal.  The  fourth  digit  contains  only  the  metacarpal.  Of  the 
left  hand  there  are  portions  of  3  digits  preserved,  including  3  metacarpals  and  a 
phalanx,  which  in  structure  are  not  different  from  those  of  the  right  hand. 

The  ilium  of  the  left  side  is  preserved,  apparently  entire.  It  is  elongate  and  cylin- 
drical, its  upper  end  adjoining  the  twenty-eighth  vertebra.  The  head  of  the  femur 
lies  close  to  the  lower  end  of  the  ilium,  so  that  that  element  must  have  been  suspended 
in  the  flesh,  much  as  in  modern  salamanders.  It  could  not  have  been  of  much  use  as 
a  support  for  the  body.  The  form  of  the  femur  is  not  unlike  that  described  for  the 
humerus,  save  that  its  lower  end  is  smaller  than  the  upper,  while  in  the  humerus 
the  extremities  are  of  equal  diameter.  A  portion  of  the  right  femur  is  preserved, 
extending  in  an  opposite  direction  to  the  left. 

Measurements  of  the  Type. 


Length  of  entire  specimen 64 

Length  of  portion  of  skull  preserved    6.5 

Posterior  width  of  same   7 

Width  across  orbits  11 

Long  diameter  of  orbit   3 

Transverse  diameter  of  orbit   1.75 

Interorbital  width   4.75 

Length  of  dorsal  Vertebrae  48 

Length  of  caudal  scries   11 

Length  of  anterior  dorsal  centrum   2 

Length  of  anterior  dorsal  rib   4 

Length  of  anterior  caudal  rib 1 .75 

Length  of  scapula 5 

Greatest  width  of  scapula   4.25 

Probable  length  of  interclavicle  6 

Width  of  same   3 

Mazonerpeton  costatum  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  341,  pi.  2,  fig.  3;  pi.  8,  fig.  4;  pi.  9,  fig.  2;  pi.  10.     1912. 

The  remains  on  which  the  present  species  is  based  are  inclosed  in  a  much  frac- 
tured nodule.  The  parts  of  the  animal  which  have  been  identified  are:  a  part  of 
the  skull  and  left  mandible,  two  clavicles,  a  humerus,  impressions  of  several  verte- 
brae, a  portion  of  the  dorsal  region  of  the  body  with  several  ribs,  two  portions  of  the 
caudal  region  with  several  ribs  and  unidentified  fragments.     (Plate  4,  fig.  3.) 


Length  of  clavicle  4.5 

Width  of  same   1.5 

Length  of  right  humerus   6 

Distal  width  of  same   2 

Length  of  ulna  3.25 

Distal  width  of  same   1 

Length  of  radius 3 

Width  of  carpal  space   2 

Length  of  metacarpal   1 .74 

Length  of  first  phalanx   1 .75 

Length  of  distal  phalanx  of  right  hand 35 

Number  of  chevron  rods  in  I  mm 4 

Length  of  ilium   2.25 

Length  of  femur  4 

Proximal  width  of  femur    1 .50 


64 


THE    COAL   MEASURES    AMPHIBIA   OF   NORTH   AMERICA. 


rb$Lcv 


The  animal,  from  the  shape  and  form  of  the  ribs,  is  undoubtedly  a  branchio- 
saurian,  since  short,  heavy,  straight  ribs  have  not  yet  been  found  to  be  associated 
with  other  than  branchiosaurian  structures.  It  is  placed  in  the  genus  Mazonerpelon 
on  account  of  the  structure  of  the  pectoral  elements,  the  form  of  the  humerus,  and 
the  length  of  the  tail,  all  of  which  agree  in  structure  with  Mazonerpelon  longicauda- 
tum.  The  animal  attained,  perhaps,  a  length  of  4.5  inches,  while  that  of  M.  longicau- 
datum  was  about  3  inches.  The  tail  in  the  present  species  is  very  long  and  slender, 
more  elongate  than  in  any  other  known  branchiosaurian. 

The  part  of  the  skull  preserved  is  very  unsatisfactory  and,  aside  from  the  fact 
that  it  seems  to  represent  the  under  side  of  the  left  half  of  the  skull,  little  can  be 
said.  Three  sutures  can  be  observed,  but  what  sutures  they  are  is  undetermined. 
The  left  mandible  lies  crushed  on  the  edge  of  the  skull  and  partially  obscures  what 
little  there  is  of  that  structure.  The 
slightly  curved  impression,  from 
which  the  bone  has  been  either 
broken  or  weathered,  measures  13 
mm.  in  length  by  3  mm.  in  posterior 
diameter  by  1  mm.  in  anterior  diam- 
eter. These  measurements  show  the 
element  to  have  been  slender  and 
pointed  anteriorly. 

Very  little  accurate  information 
can  be  derived  from  the  study  of 
the  vertebral  column  of  the  speci- 
men, nor  can  the  dorsal  vertebral 
formula  be  made  out,  since  only  a 
portion  of  the  length  of  that  region 
is  preserved  and  only  a  few  rather 
indefinite  impressions  can  be  dis- 
cerned. These  impressions  show  Fig.  14J.— Skeleton  of  Mazonerpelon  coslatum  Moodie.  X  1.5. 
,,  ...  -         .  j  1  •    1  Original  in  Yale  University  Museum,     dv,  dorsal  vertebra;; 

the  Vertebrae  tO  be  Short  and  higher  ch,  neutral  spines;  d,  clavicle;  cv,  caudal  vertebra:;  /,  femur: 

than  in  most  Branchiosauria.  *-  humerus;  m'  mandible;  '*•  rib=  **• « ».  verteb™. 

The  caudal  series  is  represented  by  two  sections,  one  of  which  is,  apparently, 
from  near  the  base  of  the  tail,  judging  from  the  size  of  the  caudal  ribs  preserved ; 
the  other  is  from  near  the  tip  of  the  tail,  and  shows  the  constituents  to  have  been 
long  and  slender.  Ribs  are  apparently  absent  on  this  section.  The  position  of  the 
two  caudal  sections  shows  that  when  the  animal  died  it  was  coiled  up  much  like  a 
snake,  so  that  in  the  fractured  nodule  three  sections  of  the  body  are  visible.  The 
tail  was  probably  half  as  long  again  as  the  body. 

The  ribs  throughout  the  body  are  short,  heavy,  and  straight,  with,  in  the  dorsal 
series,  a  lateral  and  a  distal  expansion  which  is  taken  as  a  distinctive  specific  char- 
acter. Judging  from  imperfect  impressions  in  the  dorsal  series,  the  ribs  were 
attached  to  a  transverse  process  of  the  centrum,  thus  agreeing  with  other  branchio- 


MOODIE 


PLATE  5 


2. 


A  reconstruction  of  the  possible  appearance  in  life  of  the  Coal  Measures 
branchiosaurian,  Eumkrerpetoii  parvum  Moodie,  a  small,  primitive 
salamander,  less  than  2  inches  in  length,  based  on  three  specimens  from 
the  Mazon  Creek  Shales.  The  lateral-line  organs  are  represented  as  dark 
bands  on  the  tail,  the  sense  organs  being,  apparently,  situated  beneath 
specialized  pigmented  scales,  to  which  is  due  the  preservation  of  the  lines. 

A  restoration,  natural  size,  of  the  branchiosaurian,  Mazonerpeton ,  based  on 
two  specimens.  The  form  of  the  animal  is  quite  salamander-like.  It  is 
shown  when  about  to  feed  on  a  specimen  of  Acanthotehon  stimpsoni,  which 
is  said  to  be  a  brackish-water  crustacean.  The  branchiosaur  and 
crustacean  may  possibly  have  inhabited  the  same  body  of  water. 


THE   AMERICAN   COAL    MEASURES    BRANCHIOSAURID.^. 


65 


saurians  in  this  respect.  The  ribs  show  a  progressive  decrease  in  length  from  the 
cervical  region  to  the  point  of  their  disappearance  on  the  tail. 

The  pectoral  girdle  is  represented  by  two  elements,  one  of  which  is  certainly 
the  right  clavicle,  and  the  other  is  possibly  the  left  clavicle,  though  its  form  is  some- 
what distorted  by  pressure.  Both  elements  are  in  the  form  of  an  elongate  spatula 
with  the  dorsal  surface  greatly  concave  and  the  inner  end  acuminate. 

The  right  humerus  is  imperfectly  preserved,  though  the  impression  allows  one  to 
gain  an  exact  idea  of  its  form.  It  lies  under  the  right  clavicle.  Its  ends  are  truncate 
with  a  contracted  shaft  and  expanded  extremities;  the  bone  was  apparently  hollow. 


in 


Fig.  15. 

A.  Impression  of  Erierpeton  branchialis  Moodie.    bb,  basibranchial ;  hyp,  hypohyals;  m,  mandible; 

</,  body  impression.     X  3. 

B.  Eumicrer peton  parvum  Moodie.    o,  anus;/,  femur;  h,  humerus;  in,  intestine;  /,  liver;  st,  stomach; 

r,  radius;  u,  ulna.     X  3.3. 

C.  Larger  specimen  of  Eumicrer  peton  parvum  Moodie.     a,  anus;  d,  dorsal  lateral  line;  h,  humerus; 

in,  intestine;  ml,  median  lateral  line;  si,  stomach.      X  2.6. 

D.  Skeleton  of  Erpetobrachium  mazonensis  Moodie.    cl,  clavicle;  h,  humerus;  r,  radius;  sc,  scapula; 

11,  ulna.    X  2. 

E.  Rib  of  Mazonerpeton  costalnm  Moodie.     X  2.5. 

Originals  in  the  Yale  University  Museum. 

In  another  nodule  (No.  804,  Yale  Museum)  there  is  a  single  bone  preserved  which 
resembles,  to  a  great  extent,  a  rib  of  the  present  species  (fig.  15,  E),  although  some- 
what larger,  and  it  has  been  provisionally  identified  as  such.  The  element  is  very 
slightly  curved,  but  shows  the  expanded  head  of  the  rib  of  this  species. 


66 


THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


Measurements  of  the  Type  of  Mazonerpeton  costatum  Moodie. 


No.  800  (777),  Yale  University  Museum:  mm- 

Length  of  portion  of  skull  preserved 14 

Length  of  right  clavicle 16 

Width  of  right  clavicle 4 

Length  of  dorsal  region  represented 30 

Length  of  cervical  rib 8 

Length  of  dorsal  rib 6.5 

Length  of  caudal  rib 3 

Length  of  caudal  portion  of  body  preserved. .  .    55 


Length  of  mandible 15 

Greatest  width 6 

Length  of  right  humerus 10 

Greatest  width  of  humerus 2 

No.  804  (332),  Yale  University  Museum: 

Length  of  rib 11 

Width  of  head  of  rib 2 

Diameter  of  shaft I 


Sparodus  sp.  (?). 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  pt.  11,  p.  643,  pi.  40,  figs.  52  to  56,  1882. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  xn,  p.  75,  1895. 

Type:    Specimen  in  the  Peter  Redpath  Museum  of  McGill  University. 

Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

The  material  on  which  the  above  determination  is  based  was  collected  in  1878 
by  Sir  J.  W.  Dawson  in  the  coal  formation  at  the  South  Joggins,  Nova  Scotia. 
Nothing  has  been  collected  since  that  date 
that  would  give  additional  information  as 
to  the  nature  of  the  form  represented.  I 
give  here  Dawson's  description  of  the 
remains : 

"In  the  coaly  matter  or  mineral  charcoal 
at  the  base  of  tree  No.  10  appeared  a  few  frag- 
ments of  an  animal  which  may  possibly  belong 
to  the  above-named  genus  of  Fritsch,  though 
I  am  by  no  means  certain  of  this  identification 
or  of  the  real  nature  of  the  animal. 

"The  skull  is  represented  by  a  fragment  of  a  maxillary  or  intermaxillary  bone,  with 
blunt  conical  teeth.  It  is  smooth  or  marked  merely  with  microscopic  dots.  There  is  also  a 
fragment  which  may  be  a  palatal  bone  studded  with  minute  teeth. 

"A  few  vertebrae  associated  with  the  above  bones  are  long  and  narrow,  with  large  zyga- 
pophyses  and  long  neural  spines.     Length  of  body  (i.e.,  of  the  vertebra)  about  3  millimeters. 

"With  these  remains  are  a  few  bony  scales  different  from  those  of  any  other  species 
found  in  these  trees,  and  more  resembling  scales  of  Ganoid  Fishes.  They  are  somewhat 
rectangular  in  form,  enameled  on  the  surface  and  beautifully  sculptured  with  waving  lines. 

"In  the  same  trunk  were  found  some  teeth  and  bones  referable  to  Hylerpcton  dawsoni, 
and  it  is  not  impossible  that  the  remains  above  referred  to  may  have  belonged  to  some  crea- 
ture devoured  by  that  animal,  and  which  would  not  otherwise  have  obtained  admission  to 
the  interior  of  an  erect  tree.  The  tree  itself  had  been  removed  by  the  sea,  all  but  a  little 
of  the  base,  and  this  was  in  a  very  unsatisfactory  state,  so  that  doubt  might  even  exist  as  to 
the  limit  between  the  deposit  in  the  interior  of  the  tree  and  that  under  its  base." 


Fig.  15a. — Type  material  of  Sparodus,  consisting  of  a, 
a  tooth  (X  25);  b,  four  of  the  smaller  teeth  (in 
maxilla?)  (X  25);  c,  three  bony  scales  (X  5);  d, 
fragment  of  a  limb  bone  (X  2);  e,  avertebra(X  2). 
(After  Dawson.) 


CHAPTER  X. 

ORDER  CAUDATA  DUMERIL,   1806.     COAL  MEASURES  TO  RECENT. 

Naked-skinned,  elongate,  tailed  salamanders,  mud-puppies,  efts,  newts,  etc. 
External  gills  present  or  absent  in  adult  condition,  but  always  present  in  young. 
Limbs  short,  with  usually  4  digits  on  hand  and  5  on  foot,  but  this  is  subject  to 
much  difference.  Limbs  never  very  stout.  Carpus  and  tarsus  cartilaginous.  Skull 
roof  without  the  postparietal,  postorbital,  and  supra  temporal.  Skull  elements 
never  ornamented  and  never  cut  by  the  lateral-line  canals.  Vertebrae  consisting 
of  a  single  element ;  ribs  short,  attached  to  an  elongate  transverse  process.  Caudal 
ribs  seldom  present.  Parietal  foramen  lacking.  No  ventral  armature.  Fresh- 
water inhabitants. 

Suborder  PROTEIDA  Cope,  1868. 

This  order  agrees  generally  with  the  Caudata,  but  presents  one  most  important 
feature  of  difference  in  the  presence  of  the  opisthotic.  It  is  this  point  which  gives 
the  Proteida  its  intermediate  position  between  the  extinct  amphibians  and  the  recent 
species,  and  seems  to  indicate  a  connecting  line  from  the  Coal  Measures  down  to 
the  present.    The  structure  of  the  hyobranchial  arches  sustains  this  view. 

The  hyoid  apparatus  differs  from  that  of  other  adult  Caudata  and  resembles 
that  of  their  larvae  in  having  three  epibranchials,  instead  of  one  only.  The  second 
basibranchial  is  also  connected  with  the  first,  which  is  not  the  case  with  the  other 
Caudata.    Three  extinct  genera  are  placed  tentatively  in  this  suborder. 

Family  COCYTINID.dE  Cope,  1875. 

Cope,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  p.  12,  1875. 

The  present  family,  as  here  defined,  includes  the  forms  whose  structure  seems 
to  ally  them  with  the  modern  salamanders.  The  character  on  which  most  depen- 
dence is  placed  is  that  of  the  branchial  apparatus,  lacking  in  Hyphasma.  The  forms 
are  all  incompletely  known  and  the  family  will  doubtless  require  revision  on  acqui- 
sition of  additional  material.    Three  genera,  each  with  a  single  species,  are: 

Cocytimts  gyrinoides  Cope.    Linton,  Ohio,  Coal  Measures.     Based  on  the  ventral  impression  of  the 

skull,  with  the  well-developed  branchial  apparatus. 
Erierpeton  brattchialis  Moodic.    Mazon  Creek,  Illinois,  shales.    Based  on  impression  of  mandibles 

and  branchial  apparatus. 
Hyphasma  Icevis  Cope.    Linton,  Ohio,  Coal  Measures.    Based  on  incomplete  and  obscure  amphibian 

body,  lacking  limbs. 

Genus  COCYTINUS  Cope,  1871. 

Cope,  Proc.  Am.  Phil.  Soc,  1871,  177. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  360,  1875. 

Type:  Cocytinus  gyrinoides  Cope. 

Vertebra?  and  ribs  osseous;  teeth  on  the  premaxillary  bone,  none  on  the  maxil- 
lary; hyoid  elements  largely  developed,  an  axialhyal  with  basihyal  on  each  side, 
closely  united  with  the  corresponding  ceratohyal,  at  the  end  of  which  is  an  element 
in  the  position  of  a  stylohyal;  haemal  or  basibranchials  3,  the  anterior  2,  each  sup- 
porting 1  pleural  branchihyal,  and  the  third  supporting  one  also,  the  first  haemal 
branchihyal  on  the  inner  side  of  the  ceratohyal,  approaching  the  median  line,  and 
with  elongate  pleural  element. 

67 


68 


THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


">x/3, 


The  reference  by  Cope  of  this  genus  to  the  Caudata  is  one  of  the  most  interesting 
facts  connected  with  the  Paleozoic  Amphibia.  He  says:  "The  present  genus  is, 
then,  to  be  referred  to  the  neighborhood  of  Amphiuma  and  Protonopsis,  but  forming 
the  type  of  another  family"  (123).  He  regards  the  branchial  apparatus  as  being 
more  fish-like  than  that  of  any  of  the  modern  genera.  It  is  possible  that  Cocytinus 
gyrinoides  was  a  larval  branchiate  and  consequently  aquatic  form.  It  should  be 
more  fully  compared  with  Erierpeton  branchialis  from  the  Mazon  Creek  shales 
when  better  known,  as  well  as  with  Ilyphasma  Icevis  from  the  Linton  locality. 

All  three  of  these  forms  are  included,  provisionally,  under  the  Cocytinidae. 

Cocytinus  gyrinoides  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  xn,  p.  177,  1871. 

Cope,  Trans.  Am.  Phil.  Soc,  p.  278,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  pp.  364-365,  pi.  xxxix,  fig.  4,  1875. 

Type:  Specimen  No.  8613  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:    Linton,  Ohio,  Coal  Measures. 

Two  specimens  of  this  interesting  amphibian  are  known,  one  of  them  fairly  com- 
plete (No.  2564,  Am.  Mus.  Nat.  Hist.).     The  type  specimen 
consists  of  the  inferior  bones  of  the  cranium  in  a  fairly  complete 
state  of  preservation,  with  the  muzzle  and  its  teeth;  also  8 
anterior  vertebrae,  with  their  short  recurved  ribs. 

The  condition  of  the  hyal  elements  in  the  type  specimen  is 
as  follows:  the  haemal  elements  of  the  first  branchial  arch  are 
partially  concealed  on  both  sides  of  the  ceratohyal.  An  expanded 
truncate  face  for  attachment  to  the  axial  element  is  visible  on 
both  sides,  but  the  body  of  the  bone  is  flat  and  presents  the 
edge  of  the  specimen. 

The  first  pleural  element  proceeds  from  just  behind  the  axial- 
hyal;  it  is  longer  than  the  other  pleural  elements.  A  slender 
bone  is  visible  extending  from  the  space  between  the  cerato- 
hyal and  mandibular  angle;  it  may,  therefore,  pertain  to  the  sus- 
pensorium  of  the  jaw  as  well  as  to  that  of  the  hyiod  arch,  or 
be  squamosal  as  well  as  stylohyal.  The  second  haemal  bone  is 
slender,  but  with  an  enlarged  axial  extremity;  that  of  the  right 
side  is  not  so  well  preserved  as  to  be  safely  determined.  The 
third  haemal  elements  are  the  smallest,  and  originate  imme- 
diately in  front  of  the  occipital  condyles  and  diverge  outwards 
and  backwards.  They  are  little  curved,  subcylindric,  and  slightly 
expanded  at  the  extremities. 

Of  the  pleural  elements  the  first  and  second  are  little  curved  and  the  first  is 
marked  by  a  pit  or  foramen  on  the  under  side  near  the  distal  end,  which  is  clearly 
visible  on  both  sides  of  the  specimen.  The  third  and  fourth  pleurals  are  more  curved 
and  the  outer  ends  slightly  expanded  and  directed  backwards. 

The  obverse  of  the  specimen  (fig.  16)  shows  that  the  anterior  axialhyal  is  wedge- 
shaped.     The  lateral  basihyals  are  massive.     The  second  hasmal  branchihyal  is 


FlG.  16. — Obverse  of  Cocy- 
tinus gyrinoides  Cope, 
from  the  Coal  Meas- 
ures of  Ohio  X  2. 
pmx,  premaxillas;  mx, 
maxilla;  m,  mandible; 
ah,  axialhyal;  h,  basal 
branchihyal ;  ch,  cer- 
atohyal; bv,  haemal 
branchihyal;  b,  bll, 
bill,  bllll,  pleural 
branchihyals. 


PI  ATE  6 


.2  *•* 
a  H 

II 

11 

-*    at 


5  .if 

"C 

80 


.2  £ 

01  u 
_  3 

a  J= 

I< 

>.a 

»     . 

v 

■     u 

2  3 


»i  1> 

3  -a 

■  *-» 

*  .a 

£  v 

3  3 

J=  O 

-  u 

/  £ 

J)  «-> 

.5  •« 


5-3 
I  9 

s  I 
B  5 

a  .<= 

if 

I    M 
5     O 

£  S  x 

C   £L    2 

si! 

S    8    2 

i>    y    g 

5  -.  «• 

w   >  z 

^     z 

s  Z  ° 

S    -  H 

i  s  g 

ial 

^    o  * 


5  •£   * 
i 


ORDER   CAUDATA    DUMERIL,    l8o6.  69 

dilated,  fan-shaped  distally,  and  supports  two  pleural  elements.  The  muzzle  pro- 
jects over  the  lower  jaw  and  was  rather  broadly  truncate.  The  premaxillary  teeth 
are  cylindric  and  6  in  number  on  each  side.  The  maxillary  bone  is  represented  by  a 
lamina  at  each  lateral  extremity  of  the  premaxillary.  The  mandibular  rami  are 
very  stout,  as  are  also  the  ceratohyals.  The  vertebrae  have  possessed  some  apophy- 
ses, apparently  keel-shaped  diapophyses.    The  ribs  are  slightly  curved. 

Measurements  of  the  Type. 

mm.  mm 

Length  of  specimen   32  Length  of  axialhyal 3 

Length  of  skull   12  Length  of  postbranchial 4 

Posterior  width  of  skull   11  Width  of  vertebra   1 

Length  of  premaxilla   5  Length  of  vertebra    3 

Length  of  mandible  10  Length  of  rib    6 

The  other  specimen  of  this  species  (fig.  i6«)  is  interesting  in  having  40  consecu- 
tive vertebrae  preserved,  and  19  pairs  of  ribs  attached  in  their  natural  relations  to 
the  skull  and  hyal  elements.  There  are  a  few  hyal  elements  preserved,  but  nothing 
is  added  to  our  previous  knowledge.    The  ribs  are  quite  as  in  the  type  specimen,  as 


Fig.  16a. — Nearly  complete  specimen  of  Cocylinus  gyrinoides  Cope,  from  the 
Coal  Measures  of  Linton,  Ohio.  Original  in  the  American  Museum  of  Nat- 
ural History.     X  0.95. 

are  also  the  vertebras.  The  animal  was  apparently  a  slender,  eel-shaped  amphibian 
comparing  favorably  with  the  modern  Amphiuma  in  this  respect.  There  are  no 
indications  of  limbs  or  limb  girdles. 

Measurements  (No.  2564,  American  Museum  of  Natural  History). 

mm. 

Length  of  entire  specimen   113 

Length  of  skull   15 

Width  of  head  posterior  15.5 

Length  of  vertebra   2 

Length  of  rib   4 

Genus  ERIERPETON  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  328,  1912. 

Type:  Erierpeton  branchialis  Moodie. 

The  generic  characters  are  found,  first  of  all,  in  the  presence  of  hyobranchial 
arches  which  indicate  its  relationship  to  the  formerly  described  Cocytinus  gyrinoides 
Cope,  from  Ohio.  The  only  other  known  extinct  genera  of  Caudata  which  possess, 
or  at  least  have  preserved,  the  hyobranchial  arches  are  the  Jurassic  Hylceobatrachus 
from  Belgium  and  Lysorophus  from  the  Permian  of  Texas.  The  present  form  is  widely 
distinct  from  both  of  these  genera  in  the  shape  of  the  mandible  and  the  form  and 
arrangement  of  the  hyobranchial  bars.  The  genus  Erierpeton  finds  its  closest  ally  in 
Cocytinus,  in  the  family  Cocytinidae,  which  possibly  belongs  in  the  order  Caudata 
and  the  suborder  Proteida  of  Cope. 


70  THE    COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

Erierpeton  branchialis  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  VI,  No.  2,  pp.  329-330,  pi.  I,  fig.  3;  pi.  2,  fig.  1,  1912. 

Type:  Specimen  No.  801  (222)  5,  Yale  University,  Museum. 
Horizon  and  locality:  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  amphibian  remains  designated  by  the  above  name  consist  of  a  distinct  man- 
dible and  some  rather  indefinite  body  impressions  (plate  3,  fig.  4).  Three  elongate 
impressions  occur  between  the  rami  of  the  mandibles  (fig.  15,  A),  which,  I  suppose, 
must  represent  hyoid  bones  belonging  to  the  hyobranchial  arches.  The  lateral  ele- 
ments are  paired  and  the  median  impression  is  straight  and  lies  between  the  paired 
impressions  of  the  hyoids.  The  paired  portions  probably  represent  the  hypohyals 
or  hypohyals  plus  the  ceratohyals,  and  the  unpaired  portion  of  the  first  basibranchial, 
according  to  the  nomenclature  of  Wiedersheim  (Comparative  Anatomy  of  Verte- 
brates, 1897,  p.  86).  If  the  impressions  have  been  correctly  interpreted  the  present 
specimen  is  of  very  great  interest,  since  it  is  the  first  evidence  we  have  of  the  hyo- 
branchial arches  in  the  Amphibia  of  Mazon  Creek,  and  the  second  in  the  Carbon- 
iferous of  North  America.  Dawson  doubtfully  identified  (216)  some  elements  of 
the  Joggins  Amphibia  as  hyoids,  but  was  uncertain  as  to  their  position.  Cope 
described  fully  the  well-developed  hyobranchial  apparatus  of  Cocytinus  gyrinoides 
(123)  from  the  Coal  Measures  of  Ohio.  Among  other  Paleozoic  Amphibia  Williston 
(614)  has  described  branchial  arches  in  the  peculiar  form  Lysorophus  tricarinalus 
Cope,  from  the  Permian  of  Texas. 

The  form  of  the  impression  of  the  mandible  in  the  present  specimen  is  unlike 
anything  known  to  the  writer  among  other  Carboniferous  or  later  Amphibia.  The 
rami  are  long,  slender,  deep,  slightly  curved,  and  pointed  anteriorly.  The  anterior 
symphysis  was  not  a  complete  sutural  union,  but  was  occupied  partly  by  cartilage 
or  other  connective  tissue. 

There  are  no  definite  traces  of  the  appendicular  skeleton.  The  traces  of  the  body 
(fig.  15,  A)  indicate  an  elongated,  rather  slender  animal,  but  further  than  this  noth- 
ing can  be  said  in  regard  to  its  structure. 

The  occurrence  of  a  typically  caudate  form  in  the  Carboniferous  is  unusual  and 
complicates  still  further  our  understanding  of  the  origin  and  relationships  of  the 
early  Amphibia. 

Measurements  of  the  Type. 

mm>  mm. 

Length  of  entire  impression   50  Width  of  basibranchial     0.75 

Length  of  mandible  along  median  line 10  Length  of  hypohyal     2.4 

Width  of  mandibular  ramus   9  Width  of  hypohyal    \  c 

Length  of  basibranchial   2.5 

Genus  HYPHASMA  Cope,  1875. 

Cope,  Proc.  Acad.  Sci.  Phil.,  p.  16,  1875. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  387,  1875. 

Type :  Hyphasma  lends  Cope. 

' '  Vertebrae  osseous,  the  posterior  dorsals,  and  probably  the  caudals,  furnished  with  fan- 
like neural  spines;  limbs  unknown-(?)  wanting.  Thoracic  shields  present.  Ventral  arma- 
ture, consisting  of  rhomboidal  scuta,  forming  packed  rows  arranged  in  chevrons,  directed 
backwards,  on  top  of  which  are  the  usual  rod-like  scales  arranged  in  packed  chevrons,  with 
the  angle  directed  forward. 


ORDER   CAUDATA   DUMERIL,    l8o6.  7 1 

' '  The  general  appearance  of  the  type  of  this  genus  is  that  of  a  Ptyonius,  but  the  ventral 
armature  is  different  from  anything  observed  in  the  known  genera  of  this  group.  The 
larger  external  scuta  are  like  those  of  the  species  of  Colosteus  (Sauropleura) ,  but  their  series 
have  a  different  direction.    The  inner  chevrons  are  those  of  many  other  genera"  (123). 

Hyphasma  laevis  Cope. 

Cope,  Proc.  Acad.  Sci.  Phil.,  p.  16,  1875, 

Cope,  Geol.  Surv.  Ohio.,  11,  pt.  11,  p.  387,  pi.  37,  fig.  4,  1875. 

Type:  Specimen  No.  9023  (in  counterpart),  American  Museum  of  Natural 
History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

"In  the  only  known  specimen  the  vertebrae  have  low  and  squarely  truncate  neural 
spines  near  the  head,  and  some  distance  anterior  to  the  tail  they  are  quite  conspicuous 
and  delicately  line-grooved.  The  body  is  slender  and  probably  limbless.  The  thoracic 
scuta  are  large  and  close  to  the  head;  the  median  is  produced  at  both  ends,  but  chiefly 
anteriorly  while  the  lateral  are  narrow;  all  are  without  sculpture  The  head  is  seen  from 
below.  The  mandibular  rami  are  not  so  slender  as  in  most  species  of  Ptyonius,  but  are 
rather  stout.  They  are  a  little  incurved  distally,  so  that  the  form  of  the  muzzle  is  somewhat 
narrowed,  but  not  produced.  The  teeth  are  not  visible.  Ten  rows  of  the  outer  layer  of 
scuta  in  0.005  m-"  (I23)- 

The  specimen  is  very  indistinctly  preserved  and  the  characters  given  by  Cope 
can  not  all  be  made  out.  It  is  puzzling  to  see  just  on  what  he  bases  his  conclusion. 
It  is  possible  that  the  specimen  is  a  poorly  preserved  Ptyonius.  The  outlines  of  the 
vertebrae  are  so  indistinct  that  I  am  uncertain  about  them.  In  certain  lights  there 
appear  to  be  regular  impressions  which  resemble  the  spines  of  the  vertebrae  of 
Ptyonius,  but  they  are  doubtful.  The  skull  appears  totally  distinct  from  any 
known  species  of  Ptyonius,  but  it  is  very  imperfect.  The  condition  of  the  pectoral 
elements  is  very  uncertain  and  I  can  not  be  sure  that  what  Cope  described  as  tho- 
racic "scuta"  are  such.  The  interclavicle,  however,  is  clearly  preserved  as  a  diamond- 
shaped  structure.  It  is  almost  smooth,  with  a  few  faint  radiating  lines  near  the  base. 
It  measures  5  mm.  in  greatest  breadth  by  8  mm.  in  length. 

MEASUREMENTS  OF  THE  TYPE. 

mm.  mm. 

Length  of  specimen  as  preserved 64  Length  of  median  thoracic  scuta   10 

Length  of  skull   15  Width  of  same   4 

Greatest  width  of  skull   8  Width  of  clavicle   2 

Width  of  body   8  Length  of  mandibular  ramus  12 

Length  of  7  cervical  vertebrae   15 


CHAPTER  XI. 

DEFINITION  OF  THE  ORDER  SAL1ENTIA,  LAURENT),   1768. 
(World-wide  distribution.) 

Naked,  tailless  Amphibia  of  compact  form,  and  with  usually  proccelous  vertebrae.  Cau- 
dal vertebrae  coalesced  into  a  slender  elongate  piece,  the  urostyle.  Two  elements  of  the  tar- 
sus ossified  and  greatly  elongated.  Development  by  metamorphosis;  gills  never  present 
in  adult.    Ilium  greatly  elongated. 

The  order  is  suggested  in  the  Coal  Measures  by  a  single  species,  known  from  a 
single  poorly  preserved  specimen  (plate  24,  fig.  1 ) .  The  form  Pelion  lyelli  Wyman  was 
the  first  known  of  the  Linton  Amphibia,  and  its  striking  frog-like  (123,  639)  appear- 
ance was  early  noticed.  There  is  no  assurance  that  the  species  belongs  with  this  order, 
but  since  a  well-developed  and  highly  specialized  frog  (480,  481)  occurs  in  the  Como 
Beds  (405)  of  Wyoming,  it  is  not  impossible  that  we  may  have  in  the  Pelion  lyelli  a 
suggestion  (460),  at  least,  of  the  ancestral  structure.  It  is  certain  that  the  frogs  have, 
in  past  ages,  had  a  much  greater  length  of  vertebral  column  than  they  possess  at  pres- 
ent, as  is  witnessed  by  the  coalescence  of  several  vertebras  to  form  the  urostyle.  It 
is  suggested  that  the  ancestral  vertebral  column  is  represented  in  Pelion. 

Family  PELIONTIDjE  Cope,  1875. 
Cope,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  p.  11,  187.5. 

The  present  family  includes  but  a  single  species,  that  of  Pelion  lyelli  Wyman, 
first  described  in  1858  (640),  from  Linton,  Ohio. 

The  family  characters  are  to  be  found  in  the  broad  and  obtusely  rounded  cra- 
nium, in  the  frog-like  scapular  arch,  the  frog-like  hind  limb,  and  in  the  form  of  the 
palate,  so  far  as  these  structures  have  been  preserved. 

It  has  been  suggested  that  the  present  form  shows  decided  affinities  with  the 
frogs  of  to-day  and  it  may  possibly  be  looked  upon  as  the  actual  ancestor  of  the  liv- 
ing frogs.  The  length  of  the  vertebral  column  would  seem  to  militate  against  such 
a  relationship,  since  it  is  well  known  that  frogs  have  had  a  short  vertebral  column 
since  the  Jurassic  (480,  481).  But  this  is  not  a  good  argument,  since  the  developing 
urostyles  of  modern  tadpoles  show  metameric  fenestrations  in  the  developing  bone 
which  doubtless  correspond  to  openings  between  the  vertebras.  The  notochord  of 
the  tail  is  segmented,  apparently  through  the  influence  of  former  vertebral  structure. 
At  any  rate,  the  suggestion  is  an  interesting  one  and,  whether  sustained  or  disproven, 
the  present  discussion  is  based  on  the  probabilities  of  the  case. 

Genus  PELION  Wyman,  1868. 

Wyman,  Am.  Jour.  Sci.  (2),  xxv,  p.  160,  1858  (Raniceps). 

Cope,  Proc.  Acad.  Nat.  Sci.  Phil.,  1868,  211  (Pelion,  suggested  in  letter  to  Cope  by  Wyman). 

Type:    Pelion  lyelli  Wyman. 

' '  The  only  specimen  of  the  species  exhibits  an  inferior  view  of  a  portion  of  the  skeleton ; 
and  the  obverse,  on  which  the  thoracic  and  abdominal  armor  could  have  been  preserved,  has 
not  come  under  my  observation.  The  specimen,  however,  does  not  exhibit  any  ribs,  although 
the  vertebrae  are  well  preserved.  As  observed  by  Professor  Wyman,  the  genus  pre- 
sents some  points  of  similarity  to  the  Anura  (Salientia) .  The  prolongation  of  the  angles  of 
the  mandible  is  of  this  character,  as  well  as  the  general  form  of  the  head.    The  bones  of  the 

72 


MOODIE 


PLATE  7 


Uvlerpeton  dawtoni  Owen.  Above:  Mandible,  teeth,  rib,  and  bones  of  anterior  extremity. 
Below:  Bones  of  pelvis  and  posterior  limb  and  bony  scales.  Nearly  natural  size.  Erect 
tree,  Coal  formation,  South  Joggins,  Nova  Scotia.  Photograph  by  Dawson,  published 
through  the  courtesy  of  Dr.  Arthur  Willey.  Original  specimens  in  Peter  Redpath  Museum 
of  McGill  University. 


DEFINITION   OF   THE   ORDER    SALIENTIA,    LAURENTI,    1 768.  73 

forearm  may  be  united  as  in  the  frogs,  and  the  length  and  curvature  of  the  femur  are  seen 
among  these  animals  rather  than  the  Salamanders.  The  form  of  the  femur  is  different  from 
that  of  Amphibamus  grandiceps  Cope,  which  also  differs  in  the  presence  of  dermal  scales 
and  ventral  scutellae."     (123.) 

Pelion  lyelli  Wyman. 

Wyman,  Am.  Jour.  Sci.  (2),  xxv,  p.  160,  1858. 

Cope,  Proc.  Acad.  Nat.  Sci.  Phil.,  1868,  p.  211. 

Cope,  GeoL  Surv.  Ohio,  11,  pt.  11,  p.  390,  pi.  xxvi,  fig.  1,  1875. 

Moodie,  Pop.  Sci.  Monthly,  lxxii,  p.  562,  fig.  1,  1908. 

Type:  Specimen  No.  7909  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.     (Plate  24,  fig.  1.) 

This  was  the  first  species  described  from  the  Linton,  Ohio,  deposits.  It  was  made 
known  by  Dr.  Wyman  in  1857  at  the  meeting  of  the  American  Association  for  the 
Advancement  of  Science  for  that  year.  The  species  was  subsequently  studied  by  Cope. 
He  merely  confirmed  Wyman's  observations.  The  following  description  is  based  on 
the  descriptions  of  Cope  and  Wyman  and  on  my  own  study  of  the  type  specimen. 

This  is  the  most  frog-like,  in  appearance  at  least,  of  all  the  Amphibia  which 
have  so  far  been  discovered  in  the  Carboniferous.  The  skull  especially  has  a  shape 
which  is  strikingly  frog-like,  and  the  long  hind  limbs  lend  further  likeness  to  the 
tailless  forms.  Pelion  may  have  been  a  jumping  creature,  if  we  may  judge  from  its 
long  hind  legs.  Wyman  and  Cope  have  both  called  attention  to  the  frog-like  appear- 
ance of  the  specimen,  and  this  is  apparent  at  the  first  glance.  It  is  probable  that 
the  resemblance  has  some  significance  as  to  the  ancestry  of  the  Salientia,  and  it 
may  indicate  the  first  step  in  the  origin  of  the  tailless  Amphibia.  It  is  possible  that 
the  frogs  began  to  be  separated  from  the  other  Amphibia  during  the  Carboniferous. 
The  first  frogs  we  know  are  from  the  Jurassic,  where  they  are  well-developed 
ranids.  If  Pelion  be  a  frog  ancestor,  then  the  history  of  the  group  from  the  Coal 
Measures  to  Jurassic  is  an  unknown  story. 

The  specimen  is  preserved  on  its  back  and  it  is  thus  impossible  to  tell  as  to  the 
structure  of  the  skull.  Cope  was  of  the  opinion  that  the  depressed  areas  on  the 
sides  of  the  elongate  parasphenoid  were  the  orbits,  and  if  so  the  resemblance  to  the 
frogs  is  much  more  striking.  In  the  frogs  there  is  a  strong  process  from  the  ptery- 
goid which  projects  inward  to  meet  a  corresponding  process  from  the  parasphenoid. 
This  forms  a  heavy  rod  behind  the  palatine  vacuity.  There  is  a  heavy  rod  repre- 
sented in  the  specimen  and  a  part  of  it  is  certainly  the  external  process  of  the  para- 
sphenoid, but  whether  it  is  to  be  interpreted  as  in  the  frog  is  an  open  question.  The 
outline  of  the  cranium  is  partially  obscured  by  the  mandibles,  but  the  anterior  part 
is  represented  by  a  raised  line,  as  shown  in  figure  17.  In  the  anterior  part  of  this 
space  there  are  two  ridges  which  may  be  tooth  ridges.  If  they  are  teeth  there  is  a 
great  similarity  to  the  premaxillary  and  vomerine  teeth  of  Necturus,  since  the 
ridges  are  widely  separated  at  the  median  line  and  approximated  distally,  as  they 
are  in  Necturus.  The  mandible  is  preserved  entire  and  its  form  is  strikingly  frog- 
like. Its  posterior  angles  project  over  the  quadrate  area  and  seem  to  have  had  an 
upturned  projection  such  as  is  found  in  the  mandibles  of  the  Crocodilia. 

There  are  impressions  of  20  vertebras  preserved,  and  they  cover  a  little  more 
than  half  of  the  presacral  region.    There  may  have  been  28  to  30  presacrals.    The 


74 


THE    COAL    MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 


vertebrae,  as  preserved,  are  somewhat  quadrate  in  outline  and  constricted  at  the  mid- 
dle, as  though  they  were  of  the  typical  microsaurian  type.     No  ribs  are  preserved. 

There  is  an  impression  anterior  to  the  right  humerus  which  may  represent  a 
part  of  the  pectoral  girdle,  but  its  form  is  so  obscure  that  it  can  not  be  determined. 
The  pelvic  girdle  is  entirely  wanting  in  the  specimen.  Remains  of  the  fore  and  hind 
limbs  are  preserved.  The  arms  are  especially  well  pre- 
served and  consist  of  a  strong  humerus,  a  separate  radius  and 
ulna,  and  phalanges,  the  carpus  having  undoubtedly  been 
cartilaginous,  since  there  are  no  traces  of  carpal  bones. 
Wyman  has  figured  a  small  ossicle  (fig.  17)  which  might  be 
interpreted  as  a  carpal,  but  it  is  further  removed  from  the 
carpal  region  than  his  figure  shows  and  I  would  interpret  it 
as  a  fragment  of  a  phalange,  since  the  first  digit  seems  to 
be  turned  aside  over  the  vertebral  column.  The  right  hand 
is  but  imperfectly  preserved,  but  the  left  hand  is  nearly 
entire.  There  are  evidences  of  4  digits,  possibly  5.  The 
metacarpals  are  elongate  and  rather  stout.  The  phalanges 
of  the  distal  series  have  been  lost,  so  the  phalangeal  formula 
can  not  be  determined.  On  the  whole,  the  hand  has  a  very 
broad  aspect  and  is  not  at  all  slender,  as  in  the  majority 
of  the  microsaurians  from  the  Coal  Measures  of  Ohio.  It 
resembles  in  a  great  measure  the  broad  hand  of  a  toad  and 
may  thus  be  indicative  of  a  terrestrial  life.  The  humerus  is 
well  developed  and  has  pronounced  swellings,  as  though  for 
the  attachment  of  strong  muscles.  These  indications  would 
favor  the  view  of  the  animal  being  a  land  dweller. 

The  femur  and  a  part  of  the  tibia  (?)  of  the  right  side 
are  all  there  is  preserved  of  the  hind  limb.  These  elements  show  the  leg  to  have 
been  quite  long,  though  weaker  than  the  fore  limb.  The  femur  has  a  large  distal 
articular  surface.     The  fibula  is,  apparently,  absent,  though  it  may  simply  be  lost. 

The  genus  Pelion  stands  alone  among  the  Carboniferous  Amphibia.  The  form 
can  not  be  placed  in  the  order  Branchiosauria  on  account  of  the  well-developed 
limb  bones  and  the  large  mandible.  It  may  belong  with  the  Microsauria.  I  have 
placed  it  under  the  Salientia  in  the  hope  of  learning  more  about  the  early  relatives 
of  the  tailless  forms.  There  is  no  assurance  at  all  that  it  is  even  ancestral  to  the 
Salientia,  but  the  resemblances  are  striking. 

The  following  gives  the  measurements  of  the  type  specimens : 

Measurements  of  the  Type  Specimen.* 


Fig.  17. 


Peliim  lyclli  Wyman, 


an  amphibian  from  the  Coal 
Measures  of  Ohio,  the  sup- 
posed ancestral  salientian. 
(After  Wyman.)  X  0.75. 
pmx,  premaxilla;  pv,  palatine 
vacuity;  m,  mandible;  sc, 
scapula-coracoid ;  /;,  hume- 
rus; r-u,  radius  and  ulna;/, 
femur;  /,  tibia. 


Length  of  specimen,  as  preserved    no 

Median  length  of  the  skull    24 

Width  across  the  mandibular  angles   25 

Greatest  width  of  skull   30 

Length  of  vertebral  column  from  occiput  to 

sacral  region   80 

Length  of  left  humerus   19 

Width  of  distal  end  of  left  humerus   5.5 


Length  of  radius  and  ulna   1 1.5 

Width  of  distal  end  of  radius 2 

Length  of  digit  II,  as  preserved   16 

Ixngth  of  digit  in,  as  preserved   14 

Length  of  femur   24 

Width  of  distal  end  of  femur  4 

Length  of  tibia  18 


*The  type  specimen  was  collected  in  1857  by  Dr.  John  V.  Lauderdale,  who  presented  it  to  Dr.  J.  S.  Newberry. 


CHAPTER  XII. 

SUBCLASS   LEPOSPONDYL1A   Z1TTEL,   1887.     COAL  MEASURES  TO   PERMIAN. 

(Europe  and  North  America.) 

The  group  is  here  defined  according  to  the  English  edition  of  Zittel's  Text  Book 
of  Paleontology,  1902,  p.  125.  "Notochord  persistent  and  enclosed  in  constricted 
bony  cylinders,  hour-glass-shaped  in  longitudinal  section.  Teeth  simple,  conical, 
hollow. "  According  to  Zittel  there  are  two  families,  the  Microsauridae  and  the  Ais- 
topodidae.  The  latter  family  is  dealt  with  under  Aistopoda  (p.  76)  and  it  is  there 
shown  that  the  group  is  in  no  wise  a  valid  one.  The  former  family  is  regarded  as  an 
order  and  is  fully  entitled  to  that  rank.  As  defined  here  the  subclass  Lepospondylia 
contains  but  a  single  order,  the  Microsauria. 

Extinct,  terrestrial,  aquatic,  or  semi-aquatic  amphibians;  skull  pitted  and 
grooved  by  lateral-line  canals  and  by  sculpturing  marks,  or  the  skull  may  be  smooth; 
teeth  present  on  most  of  the  palate  bones;  exoccipitals  cartilaginous  or  calcified, 
never  completely  osseous ;  sclerotic  plates  sometimes  present ;  skull  of  various  shapes. 
Vertebra?  with  notochord  largely  persistent,  hour-glass-shaped;  neural  spines  low  or 
high,  or  absent;  ribs  intercentral  and  single-headed,  with  an  incipient  tubercle  in 
some  forms;  vertebral  column  differentiated  into  dorsal  and  caudal  series;  cervical 
series  not  clearly  defined.  Limb  bones  with  well-ossified  perichondrium,  endochon- 
drium  partly  ossified;  epiphyses  absent;  carpus  and  tarsus  (tarsus  osseous  in  two 
species)  cartilaginous;  phalanges  clawed  or  not;  digits  4  in  hand  and  5  in  foot. 
Pubis  sometimes  calcified  but  never  osseous. 

Definition  of  the  Order  Microsauria  Dawson,  1863.       Coal  Measures 

and  Permian. 

(Europe  and  North  America.) 

Lizard-like,  sometimes  longicaudate,  stegocephalous,  lepospondylous,  ambu- 
latory or  legless  amphibians;  skull  bones  usually  sculptured  with  pits  and  grooves; 
lateral-line  canals  well  developed  on  skull  bones;  skull  with  horns  from  tabulare 
and  supratemporals  or  without  horns ;  branchiae  never  persistent ;  sclerotic  plates 
present;  orbits  usually  well  forward.  Vertebra;  hour-glass-shaped;  endochondral 
bone  weakly  developed  throughout  skeleton,  especially  in  vertebras;  notochord 
largely  persistent;  neural  spines  low  and  rudimentary  or  long,  fan-shaped,  and 
highly  ornamented.  Dorsal  series  of  vertebral  column  variable;  usually  from  22 
to  30;  tail  containing  sometimes  over  75  vertebras,  or  tail  very  short  with  2  weakly 
developed  vertebrae;  caudal  ribs  present,  in  those  forms  with  long  caudal  series 
the  distal  vertebras  sometimes  exhibiting  2  pleurocentra.  Ribs  long  and  curved, 
always  intercentral  in  position,  single-headed,  with  at  times  an  incipient  tubercle. 
Pectoral  girdle  composed  of  scapulas,  clavicles,  coracoids,  and  interclavicle. 
Pelvic  girdle  composed  of  osseous  rod-like  ilium,  plate-like  ischium;  pubis  carti- 
laginous, sometimes  calcified.  Limbs  present  or  wanting  or  weakly  developed, 
sometimes  present  in  front  and  wanting  behind.  Radius  and  ulna  and  tibia  and 
fibula  free ;  carpus  and  tarsus  usually  cartilaginous ;  digits  4  in  hand  and  5  in  foot, 
terminal  phalanges  sometimes  clawed.  Phalangeal  formula  for  the  hand  2-2-3-2, 
for  the  foot  2-2-3-4-3.  Abdomen  covered  with  dermal  armature  composed  of 
osseous  or  corneous  rods  or  scutes ;  overlapping  scales,  fish-like  in  appearance,  some- 
times present  over  the  entire  body ;  body  also  covered  with  lizard-like  scales  or  naked. 

75 


76  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

The  order  Microsauria  was  established  by  Sir  William  Dawson  in  1863  (208)  as 
a  family  of  "reptiles"  for  the  reception  of  the  genera  Hylonomus,  Hylerpeton,  Smi- 
lerpeton,  and  Fritschia.  Hylonomus  lyelli  is  the  type  species  of  the  order.  Dawson 
(216,  p.  635)  says  of  the  species  Hylonomus  lyelli  Dawson:  "It  is  the  type  of  the 
genus  Hylonomus  and  of  the  family  Microsauria."  The  forms  which  have  been 
referred  to  the  genus  Hylonomus,  and  hence  to  the  order  Microsauria,  from  the 
deposits  of  Europe  are  discussed  under  Hylonomus. 

The  Microsauria  have  been  regarded  by  the  writer  and  others  as  being  ancestral 
in  a  sense  to  some  of  the  later  reptiles  (469) ,  but  there  seem  to  be  insuperable  obsta- 
cles in  the  way  of  a  direct  derivation  of  the  reptiles  from  the  Microsauria.  One  of 
these  obstacles  seems  to  be  found  in  the  structure  of  the  hand.  In  all  Microsauria, 
so  far  as  is  known,  there  is  no  evidence  of  more  than  4  digits  in  the  hand,  while  no 
true  primitive  reptile  possessed  less  than  5.  The  carpus  of  all  true  reptiles  is  osseous, 
while  that  of  the  Microsauria  is  merely  cartilaginous.  It  is  possible  that  the  Micro- 
sauria stand  in  some  such  ancestral  relation  to  the  later  reptiles  as  the  Crossoptery- 
gia  (489ft)  do  to  the  Amphibia.  The  Microsauria  had  undergone  adaptive  modifica- 
tions as  to  structure  and  habit,  so  that  they  have  paralleled  many  of  the  groups  of 
reptiles,  but  their  structure  is  quite  different.  The  evidence,  as  far  as  we  can  see 
now,  points  to  a  close  genetic  relation  between  the  reptiles  and  the  Microsauria,  but 
that  this  relation  is  ancestral  I,  for  one,  am  not  ready  to  say. 

The  Group  Aistopoda  Miall,  1873,  is  untenable. 

The  group  Aistopoda  was  established  in  1873  as  section  ix  by  the  Committee  of 
the  British  Association  for  the  Advancement  of  Science,  in  their  "Tabular  View  of 
the  Classification  of  the  Labyrinthodonts."  L.  C.  Miall  (449,  450)  was  the  secre- 
tary of  the  committee,  and  the  report  was  published  in  two  parts.  Two  genera 
were  at  that  time  attributed  to  the  Aistopoda,  Ophiderpeton  and  Dolichosoma,  both 
described  by  Huxley  from  the  Coal  Measures  of  Kilkenny,  Ireland. 

Fritsch  (251,  pp.  107-126)  in  1883  refers  to  the  group  as  "Familie"  and  describes 
4  genera  and  9  species  as  belonging  to  the  group.  Zittel  (642,  p.  383)  refers  to  the 
group  as  "Familie"  and  places  5  genera  in  it.  Smith- Woodward  (Vertebrate  Pale- 
ontology, 1898,  p.  129)  refers  the  Aistopoda  to  a  suborder.  In  Eastman's  trans- 
lation of  Zittel's  Paleontology  the  group  is  called  a  family,  "  Aistopodidas. "  Lydek- 
ker  (393,  p.  205)  regards  the  group  as  a  suborder.  The  writer  (469)  refers  the 
Aistopoda  to  an  order.  The  group  Aistopoda  has  been  adopted  by  practically  all 
paleontologists  and  zoologists  who  have  had  occasion  to  refer  to  these  animals. 

Lydekker  (393)  in  1890  defined  the  group  as  follows: 

"Body  long  and  snake-like,  without  limbs,  and  apparently  without  pectoral  or  pelvic 
girdles.  Vertebrae  with  elongated  centra  and  aborted  neural  spines.  Ribs  slender,  and 
barbed  like  those  of  fishes.  Teeth  smooth,  without  plications  of  the  dentine.  External 
gills  probably  persistent." 

If  now  we  take  up  a  consideration  of  each  of  the  characters  mentioned  by  Lydek- 
ker we  find  that  the  first  one  holds  good  for  all  examples  of  the  group.  The  second 
character,  "without  limbs, "  is  not  good.  Species  of  QLstocephalus ,  Ptyonius,  Molgo- 
phis  all  possess  limbs;  and  doubtless  Ophiderpeton  will  be  found  to  possess  limbs 


PLATE  8 


Fritschia  (iirtidentata  Dawson.  Above:  Bones  of  skull  and  anterior  extremity,  and  bony  rods  of  belly. 
Below:  Bones  of  pelvis  and  posterior  extremity.  Nearly  natural  size.  Erect  tree,  Coal  formation, 
South  Joggins,  Nova  Scotia.  Photograph  by  Dawson,  published  through  the  courtesy  of  Dr.  Arthur 
Willey.     Original  specimen  in  the  Peter  Redpath  Museum  of  McGill  University. 


SUBCLASS   LEPOSPONDYLIA   ZITTEL,    1 887.  77 

also,  since  it  has  a  well-developed  pectoral  girdle.  The  limbs  in  all  of  these  genera 
are  small.  The  third  character,  "apparently  without  pectoral  and  pelvic  girdles,"  is 
not  at  all  a  good  character,  since  nearly  every  specimen  of  some  species  and  almost 
all  species  show  evidences  of  pectoral  girdles  and  a  few  exhibit  pelvic  girdles.  The 
fourth  character,  "vertebrae  with  elongated  centra  and  aborted  neural  spines,"  is 
not  a  good  distinguishing  character,  since  Amphibamns ,  an  undoubted  microsaurian, 
possesses  the  same  vertebral  characters.  "Ribs  slender,  and  barbed  like  those  of 
fishes  "  is  a  character  which  is  common  to  several  widely  distinct  genera.  All  Micro- 
sauria  possess  long,  slender  ribs,  and  the  barbed  condition  is  one  which  is  possessed 
by  only  a  few,  Thyrsidium,  Ophiderpeton,  etc.,  the  so-called  "barb"  being  merely  a 
highly  exaggerated  tuberculum.  The  teeth  of  nearly  all  Microsauria  are  smooth, 
so  that  the  character  "teeth  smooth"  is  not  a  good  one  for  a  group  definition.  It 
has  not  been  possible  to  examine  any  of  the  American  specimens  for  the  plication 
of  the  dentine,  since  the  forms  are  so  rare  and  the  fossils  very  fragile.  The  last 
character,  "external  gills  probably  persistent, "  is  certainly  not  true  for  the  American 
species,  and  the  evidence  for  the  European  species  is  negative.  Fritsch  described 
and  figured  (Fauna  der  Gaskohle,  Bd.  I,  1883,  p.  114,  Tafeln  18  and  23)  structures 
which  he  regarded  as  supporting  structures  for  the  external  branchiae.  He  says  in 
regard  to  these  structures: 

"  Bei  dem  Umstande,  dass  sie  von  der  Kiemengegend  aus  sich  buschelformig  verbreiten 
und  man  ihren  Contakt  mit  einer  Art  von  Branchiae  constatiren  kann,  zweifle  ich  nicht 
daran,  dass  diese  Stabchen  dem  Kiemenapparate  angehoren.  Bedenklich  ist  nur  ihre 
grosse  Zahl  und  das  Vorkommen  bis  zum  i6ten  Wirbel  und  ich  erwog  die  Moglichkeit,  dass 
diese  Stabchen  einem  zarten  Bauchpanzer  angehoren  konnten.  Da  aber  weiter  im  Verlaufe 
der  ganzen  Wirbelsaule  nichts  Aehnliches  vorkommt,  so  ist  man  gezwungen  anzunehmen, 
dass  Dolichosoma  sehr  grosse  lange  Kiemenbuschel  besessen  haben  muss." 

John  Samuel  Budgctt  (79,  p.  162),  in  his  discussion  of  the  "Structure  of  the  Larval 
Polyp terus, "  refers  to  the  above-described  specimen  of  "aistopodous  Stegocephali, " 
i.e.,  Dolichosoma  longissimum  Fritsch,  and  calls  especial  attention  to  the  similarity 
of  the  external  rod  of  segmented  cartilage  on  the  hyomandibular  of  Polypterus  to 
this  structure,  to  which  Fritsch  has  assigned  a  branchiate  nature  in  Dolichosoma. 
There  is  no  doubt  in  the  mind  of  the  present  writer,  however,  that  the  rod  of  carti- 
lage, referred  by  Fritsch  to  the  gills,  can  be  other  than  scutellate  rods  of  the  ventral 
armature,  these  rods  belonging  to  the  armature  of  the  breast  or  throat.  The  evidence 
for  this  conclusion  is  furnished  by  Fritsch  himself  ( Fauna  der  Gaskohle ,  Bd .  1 ,  plate  1 8 , 
fig.  11),  where  all  may  read  in  the  figure  of  the  specimen  the  facts  of  the  case.  There 
is  quite  evidently  no  justification  for  Fritsch's  conclusion  of  the  branchiate  nature  of 
Dolichosoma.  There  is  no  evidence  of  any  gill-like  structure  in  the  American  snake- 
like amphibians  of  the  Coal  Measures. 

Reviewing,  then,  the  characters  of  the  group  which  have  been  assigned  by  vari- 
ous observers,  it  will  be  seen  that  there  is  but  a  single  character  which  holds  good: 
"body  long  and  snake-like."  This  is  totally  insufficient  for  the  retention  of  the 
group.  I  therefore  propose  to  abolish  the  group  entirely  from  zoological  classifica- 
tion. It  is  not  even  a  family.  It  will,  however,  be  convenient  to  refer  to  the  snake- 
like forms  as  "aistopodous." 


CHAPTER  XIII. 

THE    MICROSAURIAN    FAMILY    HYLONOMID/C,  FROM   THE    COAL  MEASURES  OF 

NOVA  SCOTIA. 

Family  HYLONOMID^  Fritsch,  1883. 

Fritsch,  Fauna  der  Gaskohle  und  der  Kalksteine  der  Perm  formations  Bohmens,  Bd.  1,  p.  159,  1883. 
Lydekker,  Cat.  Fossil  Reptilia  and  Amphibia,  iv,  p.  201,  1890. 

The  following  characterizations  of  the  family  are  those  given  by  Lydekker  (393, 
p.  201)  based  on  Fritsch  (251):  Body  slender  and  lizard -like;  skull  narrow,  with 
smooth  or  faintly  sculptured  bones ;  neural  spines  of  vertebras  well  developed,  and 
long,  slender  ribs.  Teeth  smooth,  or  with  grooved  summits.  The  whole  body 
covered  with  sculptured  scutes.    Internal  gills  may  be  developed. 

Fritsch  (251,  Bd.  1,  p.  159)  gives  the  following  in  his  original  description: 

"  Stegocephali  von  Baue  schlanker  Eidechsen  mit  schlanken  langen  Rippen.  Wirbel 
amphicoel  mit  stark  entwickelten  oberen  Dornfortsatzen.  Schadelknochen  glatt  oder 
schwach  verziert.  Schuppen  gross,  verziert,  den  ganzen  Korper  deckend.  Zahne  glatt  oder 
mit  verzierte  Spitze.  Kiemenbogen  bei  einigen  angedeutet.  Mittlere  Kehlbrustplatte 
unbekannt.     Coracoidea  ahnlich  wie  bei  Branchiosaurus  schlank,  winkelig  gebogen." 

Fritsch  includes  the  following  genera  in  the  family : 

Hylonomus,  Hyloplesion,  Smilerpeton,  Seclcya,  Orthocosta,  and  Ricnodon.  The  family  includes 
the  following  species: 

Hylonomus  lyelli  Dawson,  Nova  Scotia.  Hylonomus  wildi  Woodward,  England. 

latidens  Dawson,  Nova  Scotia.  Smilerpeton  aciedentatum  Dawson,  Nova  Scotia. 

multidens  Dawson,  Nova  Scotia.  Hylerpcton  dawsonii  Owen,  Nova  Scotia. 
wymani  Dawson,  Nova  Scotia.  intermedium  Dawson,  Nova  Scotia. 

iritschii  Gein.  and  Deichm.,  Saxony.  longidentatum  Dawson,  Nova  Scotia. 

geinilzi  Credner,  Saxony.  Fritschia  curtidentata  Dawson,  Nova  Scotia. 
(?)  pictus  Fritsch,  Bohemia. 

Genus  HYLONOMUS  Dawson. 

Dawson,  Quart.  Jour.  Geol.  Soc.  London,  xvi,  p.  274.  figs.  14-18,  i860 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  44,  18(13. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  634. 

Credner,  Zcit.  d.  dcutsch.  geol.  Gesell.,  1890  (ix  Thcil,  die  Stegocephalen  und  Saurier). 

Type :  Hylonomus  lyelli  Dawson. 

The  genus  Hylonomus  is  a  very  important  one  from  a  taxonomic  viewpoint, 
since  it  was  regarded  by  Dawson  (216,  p.  635)  as  the  typical  genus  of  the  order  Micro- 
sauria,  the  most  abundant  group  in  the  Carboniferous.  Unfortunately  the  species 
of  the  genus  Hylonomus  are  known  only  from  fragmentary  remains.  I  have  re- 
produced in  plate  9  Dawson's  figures  of  the  remains  of  Hylonomus  as  published  by 
him  in  1891. 

Dawson  (216)  gave,  in  1882,  the  following  definition  of  the  genus  Hylonomus: 
"Form  lizard-like,  with  the  posterior  limbs  somewhat  large  in  proportion  to  the 
anterior.  Size,  small.  Mandibular  and  maxillary  teeth  numerous,  small,  conical, 
pointed.    Palatal  teeth  minute.    Abdominal  scales  oval. " 

78 


MOOOti 


PLATE  9 


Hylonomus  lyelii  Dawson.  1,  maxillse  and  skull  bones;  la,  sternal  bones;  2,  mandible;  3,  humerus,  ribs,  and 
vertebra;  4,  posterior  limb;  5,  pelvis;  6,  caudal  vertebrae.  Nearly  natural  size.  Erect  tree,  Coal 
formation,  South  Joggins,  Nova  Scotia.  Photograph  by  Dawson,  published  through  the  courtesy  of 
Dr.  Arthur  Willey.     Original  in  the  British  Museum. 


THE   MICROSAURIAN   FAMILY   HYLONOMID^.  79 

Credner  (186),  Fritsch  (251,  Bd.  1,  p.  89,  Taf.  12,  figs.  1,  4,  15),  and  Woodward 
(629)  have  referred  remains  of  Microsauria  discovered  in  the  Coal  Measures  or 
lower  Permian  deposits  of  Saxony,  Bohemia,  and  Lancashire,  England,  to  the  genus 
Hylonomus.  There  is  much  uncertainty  as  to  the  validity  of  these  references,  due 
to  the  uncertain  nature  of  the  type  of  Hylonomus.  There  are  4  American  species 
of  the  genus:  Hylonomus  lalidens  Dawson,  II.  lyelli  Dawson,  II.  multidens  Dawson, 
and  //.  wymani  Dawson.  All  the  species  are  from  the  Coal  Beds  at  the  South 
Joggins,  Nova  Scotia. 

Hylonomus  lyelli  Dawson. 

Dawson,  Quart.  Jour.  Geol.  Soc.  London,  xvi,  p.  274,  figs.  14  to  18,  i860. 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  44,  1863. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  635,  pi.  39,  figs.  1  to  14  and  27. 

Dawson,  Acadian  Geology,  3d  ed.,  1880,  p.  370. 

Type:  Specimens  Nos.  R  443  to  445  in  the  British  Museum  (393,  pt.  iv,  p.  223). 
Horizon  and  locality :  Coal  formation  of  the  South  Joggins,  Nova  Scotia. 
This  species  is  by  far  the  most  abundant  (plate  9)  in  the  erect  trees  examined  by 
Dawson.     Its  characters  Dawson  (216)  defines  as  follows: 

' '  General  form  lizard-like,  with  the  hind  limbs  rather  larger  than  the  fore  limbs.  Length 
when  mature,  5  to  6  inches. 

"Head  somewhat  elongate;  bones  of  skull  smooth  or  with  microscopic  striae,  perfectly 
united,  except  at  the  parietal  foramen.  Occipital  condyle  double,  and  apparently  bony. 
Teeth  simple,  conical,  numerous,  about  forty  in  each  mandible,  and  nearly  equal,  except  that 
a  few  of  the  anterior  ones  are  rather  larger  than  the  others.  The  teeth  are  anchylosed  to 
the  jaw  in  a  furrow  protected  by  an  external  bony  plate. 

' '  Vertebras  with  cylindrical  bodies,  slightly  concave  at  the  ends.  When  partly  exfoliated 
they  appear  hour-glass-shaped,  in  consequence  of  the  internal  cartilage  having  the  form  of 
two  cones  attached  by  their  apices.  Zygapophyses  conspicuous  above ;  neural  arches  united 
to  the  bodies  of  the  vertebrae,  and  with  broad  neural  spines.  Dorsal  vertebrae  with  strong 
lateral  processes.  Caudal  vertebrae  apparently  simple  and  cylindrical.  Number  of  verte- 
brae in  neck  and  trunk  about  thirty. 

"Ribs  long  and  curved,  with  capitulum  and  tuberculum,  cartilaginous  within. 

"Anterior  limb  slender,  humerus  with  distinct  keel;  radius  and  ulna  separate;  toes  four 
or  five. 

"Posterior  limb  with  well-developed  femur;  tibia  and  fibula  shorter,  separate;  toes 
five,  somewhat  long  and  slender. 

"Pelvis  large,  composed  of  ilium  and  ischium." 

Interclavicle  and  numerous  scutellse  are  present.  Upper  surface  protected  with 
imbricated  horny  scales.  In  front  two  rows  of  horny  tubercles  and  plates,  with 
epaulettes  composed  of  bristle-like  fibers  projecting  from  the  skin. 

The  animal  possibly  fed  on  insects,  as  is  indicated  by  the  coprolitic  matter  asso- 
ciated with  the  remains  of  the  species. 

The  following  measurements  are  given  by  Dawson  for  the  largest  individual 
discovered: 

Length  of  head about  cm.  2  Length  of  rib cm.  1.3 

Length  of  neck "       "  1.3  Length  of  humerus "  1.4 

Length  of  trunk    "        "  7  Length  of  femur "  1.8 

Length  of  posterior  limb  to  heel cm.  3  Length  of  tibia "  1.2 

Length  of  mandible "  1.8  Length  of  body  of  vertebra mm.  2.5 

Teeth,  5  in  1  mm. 


80  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

Hylonomus  latidens  Dawson. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  ii,  p.  637,  pi.  39,  figs.  18-22. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  p.  74. 

Type:  Specimen  No.  3061-1,  Peter  Redpath  Museum,  McGill  University.  The 
British  Museum  (393,  pt.  iv,  p.  224)  also  has  a  specimen,  No.  R  447. 

Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

Fragments  of  3  specimens  from  3  trees  represent  this  species  (plate  10).  It  seems 
to  have  been  of  stouter  build  than  II.  lyelli,  with  the  limbs  shorter  in  proportion.  Its 
generic  affinities  are  somewhat  doubtful,  as  it  presents  in  some  respects  characters 
intermediate  between  Hylonomus  and  Hylerpeton . 

Mandibular  and  maxillary  teeth  broadly  conical,  about  20  in  each  mandible— 
3  in  1  mm. ;  anterior  mandibular  teeth  somewhat  larger  than  the  others,  and  bent  or 
hooked.  Vomer  or  palate  with  minute  teeth.  Thoracic  plate  large.  Scales  of  abdo- 
men oval,  but  somewhat  narrow,  and  tending  to  be  oat-shaped. 

Length  of  mandible  (imperfect)    mm.  9         Length  of  tibia  (?)    mm.  5 

Length  of  humerus   mm.  7         Length  of  thoracic  plate   cm.     I 

Length  of  vertebra   mm.  2  Length  of  six  caudal  vertebra;   mm.  8 

Hylonomus  multidens  Dawson. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.- 11,  p.  637,  pi.  39,  figs.  23-26. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  p.  74. 

Type:  Specimen  No.  3061-2,  Peter  Redpath  Museum,  McGill  University. 

Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

This  animal  is  known  only  by  portions  of  bones  of  the  head  and  a  few  other  frag- 
ments. The  scattered  bones  of  the  extremities  are  inseparable  from  those  of  II.  lyelli 
occurring  with  it.  As  compared  with  that  species,  the  bones  of  this  are  smoother 
and  more  delicate.  The  teeth  are  more  numerous  and  slender.  The  crushed  distal 
end  of  a  femur  or  humerus  found  near  the  skull  indicates  that  the  limbs  were  well 
developed. 

mm. 

Length  of  mandible   1 1 

Length  of  skull   15 

Length  of  femur   9 

Teeth,  5  to  6  in  1  mm. 

Hylonomus  wymani  Dawson. 

Dawson,  Quart.  Jour.  Geol.  Soc.  London,  xvi,  p.  277,  figs.  27-29,  i860. 
Dawson,  Air-breathers  of  the  Coal  Period,  p.  52,  1863. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  637,  plate  39,  figs.  15-17. 
Dawson,  Acadian  Geology,  3d  ed.,  p.  378. 

Type:  Specimen  No.  3061,  Peter  Redpath  Museum,  McGill  University.  There 
is  also  specimen  No.  R  446  in  the  British  Museum  (393,  pt.  iv,  p.  224). 

Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

As  compared  with  the  II.  lyelli  the  present  species  is  smaller  in  size,  more  elon- 
gated in  form,  had  the  teeth  less  numerous  (about  22  in  the  mandible),  and  shorter 
and  more  obtuse  in  form.    There  are  6  to  7  in  1  mm. 

This  species  is  much  more  rare  than  II.  lyelli,  but  quantities  of  minute  bones, 
probably  belonging  to  it,  occur  in  the  coprolitic  matter.    In  one  specimen  38  verte- 


PLATE    10 


Hvlonomits  latidriis  Dawson.  Skull,  portion  of  skeleton,  foot,  scapula, 
sternal  bones,  humerus,  and  rib,  believed  to  belong  to  this  species.  Erect 
tree,  Coal  formation,  Nova  Scotia.  Nearly  natural  size.  Photograph 
by  Dawson,  published  through  the  courtesy  of  Dr.  Arthur  Willey. 
Original  in  the  Peter  Redpath  Museum  of  McGill  University. 


THE    MICROSAURIAN   FAMILY   HYLONOMIOE.  8l 

brae  of  this  species  were  found  partially  associated,  indicating  a  long,  slender  body. 
The  body  is  covered  with  scales  and  ventral  scutellae  are  present.  Dawson  ques- 
tions whether  this  species  may  not  be  the  young  of  H.  lyelli. 

mm.  mm. 

Length  of  skull    8  Length  of  femur  6 

Length  of  mandible   5  Length  of  humerus  5 

Length  of  rib   5.5 

Genus  SMILERPETON  Dawson. 

Dawson,  Phil.  Trans.^Roy.  Soc,  London,  1882,  pt.  11,  p.  634. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  p.  74. 

Type :  Smilerpeton  aciedentatum  Dawson. 

The  type  species  was  originaHy  referred  to  Hylonomus,  but  further  study  induced 
Dawson  to  refer  it  to  a  new  genus.  Dawson  gives  (216)  the  following  character- 
istics of  the  genus: 

"Form  somewhat  elongated,  and  limbs  short.  Mandibular  and  maxillary  teeth  wedge- 
shaped,  with  cutting  edges.  Palatal  teeth  numerous,  some  of  them  large.  Abdominal 
scales  oval.  A  single  species  is  known,  S.  aciedentatum,  from  the  Coal  Measures  at  the  South 
Joggins,  Nova  Scotia." 

Smilerpeton  aciedentatum  Dawson. 

Dawson,  Quart.  Jour.  Geol.  Soc.  London,  xvi,  p.  275,  figs.  19  to  23,  i860. 

Dawson.  Air-breathers  of  the  Coal  Period,  p.  65,  1863. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  n,  p.  638,  plate  40,  figs.  28  to  45. 

Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  p.  75. 

Dawson,  Acadian  Geology,  3d  ed.,  p.  376. 

Type:  Specimen  No.  3061-3,  Peter  Redpath  Museum,  McGill  University.  The 
British  Museum  (393,  pt.  IV,  p.  224)  also  has  a  specimen,  No.  R  433. 

The  important  characteristic  (plate  12)  is  found  in  the  form  of  the  mandibular 
and  maxillary  teeth,  which  are  of  a  peculiar  wedge-shape,  being  broad  and  oval 
at  the  base  and  narrowed  to  a  longitudinal  edge  at  top.  Thus,  when  viewed  from  the 
side  they  appear  narrow  and  blunt,  but  when  the  jaw  is  broken  across,  and  they  are 
viewed  from  the  rear  or  front,  they  appear  broad  and  sharp-edged.  The  effect  of 
this  arrangement  is  that  the  jaw  is  armed  with  a  closely  placed  series  of  chisels  or 
wedges,  giving  an  almost  continuous  edge.  At  the  end  of  the  mandible  some  of  the 
teeth  are  longer  and  more  conical. 

Another  important  character  is  that  the  palatal  and  vomerine  bones  seem  to 
have  bristled  with  teeth,  mostly  of  very  small  size;  but  there  are  also  some  larger 
palatal  teeth,  of  which  some  are  sharply  pointed  and  others  blunt  with  furrowed 
points. 

The  vertebras  are  of  the  same  type  as  those  of  Hylonomus;  but  some  which  appear 
to  be  caudal  have  a  pointed  spine  above,  indicating  perhaps  a  flattened  tail.  The 
ribs  are  short  and  stout. 

The  body  seems  to  have  possessed  an  interclavicle  and  ventral  scutellae.  Above 
it  was,  apparently,  clothed  with  small  tubercles  and  horny  scales,  and  to  have  had 
cuticular  pendants  like  those  of  Dendrerpeton. 


82 


THE   COAL   MEASURES   AMPHIBIA    OF    NORTH    AMERICA. 


An  additional  species  of  this  genus  was  apparently  indicated  by  some  fragmen- 
tary remains,  but  Dawson  thought  best  not  to  describe  them  as  such,  since  they 
might  indicate  only  a  young  individual  of  the  present  species. 


Length  of  mandible   

Length  of  femur 

Length  of  humerus  (?) 

Length  of  vertebra   

Length  of  rib   

There  are  5  teeth  in  2  mm. 


1.5  cm. 
1.5  cm. 
1.3  cm. 

3-5  mm- 
1      cm. 


1  l/tfN 


c; 


Fig.  18. — Skeletal  elements  of  Smilcrpeton  aciedentatum  Dawson,  from  the  Coal  Measures 
of  Nova  Scotia.  (After  Dawson.)  a,  shaft  of  femur?  X  2;  6,  intermaxillary _and  teeth, 
X  25;  c,  sections  of  teeth,  X  25;  d  and  e,  palatal  teeth,  X  25;/,  femur,  X  2;  g,  rib, 
X  2;  h,  palate;  i,  caudal   vertebrae;  j,  long  palatal  tooth,  X  25;  k,  bony  scale. 

Genus  HYLERPETON  Owen. 

Owen,  Quart.  Jour.  Geol.  Soc.  London,  xvm,  p.  241,  1862. 

Dawson,  Amcr.  Jour.  Sci.  (3),  xn,  p.  443,  1876. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  634. 

Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  p.  74. 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  55,  pi.  vi,  figs.  32-46,  1863. 

Body  stout,  with  strong  limbs.  Mandibular  and  maxillary  teeth  strong, 
not  numerous,  grooved  at  apex.  Palatal  teeth  numerous,  and  some  of  them  large. 
Thoracic  plate  broad.    Abdominal  scales  pointed  or  oat-shaped. 

Type:  Hylerpeton  dawsoni  Owen. 

Hylerpeton  dawsoni  Owen. 

Owen,  Quart.  Jour.  Geol.  Soc,  xvm,  p.  241. 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  55,  1863. 

Dawson,  Acadian  Geology,  p.  380. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  n,  p.  639,  pi.  41,  figs.  62-85. 

Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1894,  xn,  p.  74. 

Type:  Specimen  No.  3061-4,  Peter  Redpath  Museum,  McGill  University. 
There  are  also  specimens,  Nos.  R  441  and  442,  in  the  British  Museum  (393,  pt.  iv, 
p.  225).     (Plate  7.) 

Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

Bones  of  skull  slightly  striated,  but  not  sculptured  as  in  Dcndrerpeton.  Lower 
jaw  with  distinct  ascending  ramus  or  coronoid  process,  a  feature  not  known  in  any 
other  of  the  Nova  Scotia  fauna,  but  observed  by  Cope  in  Brachydectes.  Teeth,  12 
in  each  ramus  of  the  mandible,  bluntly  conical,  slightly  striated  at  the  apex.  Pulp- 
cavities  large  and  longitudinally  striated  at  the  sides,  though  the  teeth  are  not 


MOODII 


PLATE    11 


Hylerpeton  longidentatunt 'Dawson.     Mandible  and  other  bones.  Nearly  natural  size.     Erect 

tree,    Coal    formation,    South    Joggins,      Nova    Scotia.  Photograph    by    Dawson, 

published  through  the    courtesy   of   Dr.    Arthur   Willey.  Original   specimen   in    the 
Peter  Redpath  Museum  of  McGill  University. 


PLATE  12 


SmiUrpetOH  aciedentatum  Dawson.  Mandible,  portions  of  skull,  scales,  and  various  bones. 
Nearly  natural  size.  Erect  tree.  Coal  formation,  South  Joggins,  Nova  Scotia. 
Photograph  by  Dawson,  published  through  the  courtesy  of  Dr.  Arthur  Willey. 
Original  specimen  in  the  Peter  Redpath  Museum  of  McGill  University. 


THE    MK'KOSAURIAN   FAMILY   HYLONOMID^E.  83 

folded.  Maxilla  furnished  with  similar  teeth,  one  of  which  near  the  front  is  larger 
than  the  others.  Palatal  teeth  numerous,  small,  conical,  with  a  few  large  teeth  at 
the  sides. 

Vertebras  short,  cylindrical,  well -ossified,  with  well-developed  zygapophyses  and 
neural  spines;  ribs  strong  and  much  curved,  with  well-developed  division  of  the 
proximal  ends;  pelvis  imperfect,  but  apparently  large,  with  broad  ilium. 

Humerus  half  the  length  of  the  mandible ;  radius  half  as  long  as  humerus ;  femur 
very  large  and  stout,  nearly  as  long  as  the  mandible;  leg  bones  and  phalanges  corre- 
spondingly stout. 

The  thoracic  plate  (plate  7)  is  indicated  only  by  some  fragments.  The  abdom- 
inal scales  are  narrow  and  pointed  (oat-shaped) ,  smooth  externally  and  with  a  ridge 
at  one  side  within.    The  following  are  the  dimensions  of  the  largest  specimen: 

Length  of  mandible 4.4  cm.  Length  of  radius 1.5  cm. 

Length  of  largest  tooth 5     mm.  Length  of  vertebra 6     mm. 

Length  of  femur 3.5  cm.  Length  of  rib 3     cm. 

Length  of  tibia 2     cm.  Length  of  scales 5  to  7     mm. 

Length  of  humerus 2     cm. 

Hylerpeton  longidentatum  Dawson. 

Dawson,  Am.  Jour.  Sci.  (3),  xn,  pp.  440-447,  1876. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  640,  pi.  42,  figs.  86  to  109. 

Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1894,  xn,  p.  74. 

Type:  Specimen  No.  3061-6,  Peter  Redpath  Museum,  McGill  University. 
There  is  also  a  specimen,  No.  R  440,  in  the  British  Museum  (393,  pt.  iv,  p.  225). 
(Plate  11.) 

Horizon  and  locality :  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

Head  much  elongated,  with  the  bones  minutely  pitted,  and  with  delicate 
microscopic  stria;,  but  not  sculptured.  Mandibular  and  maxillary  teeth  long  and 
acute,  pointing  backwards,  with  the  apex  of  their  inner  sides  finely  striated;  20 
or  more  in  each  ramus  of  the  lower  jaw ;  palatal  bones  with  several  long,  slender 
teeth  and  many  minute  teeth.  The  mandibles  found  are  not  complete,  but  there 
are  indications  that  there  was  an  ascending  process  as  in  H.  dawsoni,  but  less 
developed.  The  narrowness  of  the  dentary  bone  is  caused  in  part  by  the  lower 
posterior  edge  being  bent  inward  and  by  the  posterior  end  being  broken  off  above. 

Vertebra;  short  and  stout,  and  apparently  well  ossified.  Ribs  long,  with 
double  head  and  much  curved.  Humerus  longer  than  femur,  which  is  short  and 
stout,  if  the  bone  taken  for  it  is  rightly  determined.  Abdominal  scales  narrow, 
oat-shaped ;  thoracic  plate  large,  broadly  oval. 

Measurements  of  Hylerpeton  longidentatum  Dawson. 

Length  of  mandible 4     cm.  Length  of  femur  (?) 1.2  cm. 

Length  of  vertebra 5     mm.  Length  of  tibia 8     mm. 

Length  of  rib ' 3     cm.  Length  of  mandibular  teeth 3     mm. 

Length  of  humerus i-5  cm.  6  to  7  teeth  in  1  cm. 

Hylerpeton  intermedium  Dawson. 
Dawson,  Proc.  and  Trans.  Roy.  .Soc.  Canada,  xn,  p.  75,  1895. 

Type:  Specimen  No.  3061-5,  Peter  Redpath  Museum,  McGill  University. 
Horizon  and  locality:  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 
This  species  is  known  only  by  the  mandibles  and  portions  of  the  skull,  which  are 
rather  shorter  than  those  of  adult  individuals  of  the  last  species.    The  extremity  of 


84  THE    COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

the  mandible  and  the  cranial  bones  have  the  same  slightly  waved  surface  as  in  the 
other  species.  Mandibles  3  cm.  long  and  the  teeth,  which  are  about  15  in  each 
ramus  of  the  lower  jaw,  are  simple,  with  large  pulp  cavities,  those  of  the  maxillary 
bone  slightly  enlarging  upwards,  and  intermediate  in  form  between  the  long, 
slender  teeth  of  H.  longidentatum  and  the  thick,  obtuse  teeth  of  H.  dawsoni. 

Coal  formations,  South  Joggins,  Nova  Scotia,  in  erect  tree,  discovered  by 
P.  W.  McNaughton,  1893. 

Genus  FRITSCHIA  Dawson,  1882. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  11,  p.  634. 
Lydekker,  Cat.  Fossil  Reptilia  and  Amphibia,  pt.  iv,  p.  225,  1889. 

Type:  Fritschia  curtidentata  Dawson. 

Body  lizard-like ;  limbs  large  and  well-ossified,  mandibular  and  maxillary  teeth 
conical,  grooved  at  the  apex.    Abdominal  scales  slender  and  rod-like. 

Fritschia  curtidentata  Dawson. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  1882,  pt.  n,  p.  641,  pi.  43,  figs.  1 10-128. 
Dawson,  Am.  Jour.  Sci.  (3),  xn,  p.  444,  1876. 

Type:  Specimen  No.  3061-7,  Peter  Redpath  Museum,  McGill  University. 
There  is  also  specimen  No.  R  449,  in  the  British  Museum  (393,  pt.  iv,  p.  225). 

Horizon  and  locality :  Coal  formation  at  the  South  Joggins,  Nova  Scotia. 

Represented  by  2  specimens  (plate  8).  Bones  of  the  head  very  smooth,  only 
a  few  microscopic  punctures.  Teeth  conical,  somewhat  obtuse,  striated  at  the  inner 
side  of  the  apices;  there  are  about  30  in  each  ramus  of  the  mandible,  and  about  27 
in  the  maxillary  bone.  Teeth  implanted  in  a  furrow.  Vertebrae  short  and  well 
ossified,  3  in  1  cm.  Ribs  strong,  curved,  about  1  cm.  in  length.  Limbs  robust, 
the  bones  better  ossified  than  in  any  other  known  species  from  Nova  Scotia.  Toes 
of  foot  probably  5,  central  ones  long  and  slender.  Interclavicle  of  moderate  size 
and  somewhat  rounded.    Ventral  scutellas  needle-like. 

Length  of  mandible  (imperfect) 2.1  cm.  Length  of  femur 2.4  cm. 

Length  of  maxilla 2     cm.  Length  of  radius  and  ulna 1     cm. 

Length  of  rib 1     cm.  Length  of  toe  in  foot 7     mm. 

Length  of  humerus 2     cm.  8  teeth  in  5  mm. 


CHAPTER  XIV. 

THE  MICROSAURIAN  FAMILY  TUD1TANID/C,  FROM  THE  COAL  MEASURES 
OF  OHIO  AND  PENNSYLVANIA. 

Family  TUDITANIDiE  Cope,  1875. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  357,  1875. 

Lizard-like  microsaurians ;  cranial  elements  strongly  sculptured  with  pits  or 
grooves  or  almost  smooth,  with  weak  punctulations.  Orbits  usually  well  forward; 
squamosal  sometimes  excluded  from  the  parietal ;  skull  hornless ;  teeth  pleurodont, 
conical  and  sharp,  smooth  or  slightly  plicate;  clavicle  of  a  triangular  shape,  which 
is  characteristic  of  all  the  species;  vertebrae  well  developed  and  phyllospondylous, 
the  osseous  portion  being  merely  a  hollow  cylinder,  hour-glass-shaped ;  ribs  curved, 
long,  attenuated  and  intercentral ;  digits  clawed;  ventral  armature  absent  in  all 
but  a  single  species  and  the  association  of  the  species  is  doubtful;  tail  moderate 
in  length.    Three  genera  with  13  species  included  in  the  family.  These  species  are : 

Tuditanus  punctulatus  Cope,  Linton,  Ohio. 
brevirostris  Cope,  Linton,  Ohio. 
longipes  Cope,  Linton,  Ohio. 
minimus  Moodie,  Cannelton,  Pennsylvania. 
walcotti  Moodie,  Linton,  Ohio. 
Erpetosaurus  radiatus  Cope,  Linton,  Ohio. 
obtusus  Cope,  Linton,  Ohio. 
tabulatus  Cope,  Linton,  Ohio. 
minutus  Moodie,  Cannelton,  Pennsylvania. 
sculptilis  Moodie,  Cannelton,  Pennsylvania. 
acutirostris  Moodie,  Linton,  Ohio. 
tuberculatus  Moodie,  Linton,  Ohio. 
Odonterpeton  triangularis  Moodie,  Linton,  Ohio. 

The  association  of  these  species  in  the  one  family  is  provisional  and  will  need 
revision  on  the  acquisition  of  new  and  more  complete  material. 

Genus  TUDITANUS  Cope,  1874. 

Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  271,  1874. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  pp.  391,  1875. 

Type:  Tuditanus  punctulatus  Cope. 

The  genus  as  here  defined  is  a  somewhat  composite  group  and  it  is  quite  proba- 
ble that  some  of  the  species  here  included  will  have  to  be  removed  to  another  genus 
when  the  anatomy  of  the  forms  is  better  known.  The  species  of  the  genus  are  all 
moderately  small,  the  largest  barely  attaining  a  length  of  8  inches. 

There  are  5  species  of  Tuditanus  thus  far  known.  All  of  the  species  are  charac- 
terized by  the  possession  of  a  peculiar  triangular-shaped  clavicle  with  radiating 
grooves,  and  this  has  been  taken  as  one  of  the  distinctive  characters  of  the  genus,  as 
well  as  of  the  family.    The  structure  of  the  cranium  where  known  is  quite  uniform 

85 


86  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

among  the  different  species.    The  squamosal  is  quite  large  and  the  supratemporal 
is  not  always  closely  joined  to  the  parietal.    The  species  are: 

Tuditanus  punctulatus  Cope,  Linton,  Ohio. 
brevirostris  Cope,  Linton,  Ohio. 
longipes  Cope,  Linton,  Ohio. 
minimus  Moodie,  Cannelton,  Pennsylvania. 
walcotti  Moodie,  Linton,  Ohio. 

Tuditanus  punctulatus  Cope. 

Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  271,  1874. 

Cope,  Geol.  Surv.  Ohio,  n,  pt.  11,  p.  392,  pi.  xxxiv,  fig.  1,  1875. 

Type:  Specimen  No.  no,  American  Museum  of  Natural  History,  where  there 
is  also  specimen  No.  in. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  species,  together  with  the  form,  T.  brevirostris,  describedonp.  88,  was  used  by 
Cope  as  the  type  of  the  genus  Tuditanus.  Cope  subsequently  associated  some  rep- 
tilian remains  from  the  Linton  mines  with  the  type  of  T.  punctulatus  and  changed 
the  generic  term  to  Isodectes,  which  was  known  by  another  species,  /.  megalops  Cope, 
from  the  Permian  of  Texas.  The  remains  associated  by  Cope  with  I.  megalops 
undoubtedly  represent  a  reptilian  species  and  which  has  been  described  elsewhere 
under  the  name  Eosauravus  copei  Williston.  The  species  is  of  exceeding  interest 
because  it  is  the  oldest  known  reptile  and  places  the  range  of  the  Reptilia  down 
towards  the  base  of  the  Pennsylvanian. 

The  species  Tuditanus  punctulatus  Cope  was  founded  on  well-preserved  remains 
of  nearly  the  entire  skeleton  of  a  single  individual.  The  bones  are  represented  by 
shining  carbonaceous  matter,  and  since  both  of  the  slabs  containing  the  impression 
were  preserved,  a  great  many  characters  have  been  determined.  The  head,  I  fore- 
limb,  and  23  consecutive  vertebras  with  ribs  are  well  defined,  but  of  the  pelvis 
and  hind  limbs  nothing  is  visible. 

The  cranium  (fig.  19)  is  very  similar  to  that  of  T.  minimus,  from  the  Cannelton, 
Pennsylvania,  slates.  It  is  triangular  in  shape,  with  a  narrowed  obtuse  muzzle. 
The  orbit  of  the  left  side  is  well  defined  and  lies  well  forward.  It  is  oval  in  outline 
and  its  width  is  about  two-thirds  of  its  length.  The  nostrils  are  small  and  are 
located  well  toward  the  tip  of  the  muzzle.  The  parietal  foramen  lies  behind  the 
median  transverse  line  which  divides  the  skull  equally. 

The  cranial  elements  are  for  the  most  part  destroyed,  but  the  outlines  of  a  few 
can  be  determined.  Those  elements  which  are  preserved  are  ornamented  with  a 
sculpturing  of  minute  punctulations  which,  on  the  postfrontal,  assumes  a  radiating 
arrangement.  The  ornamentation  of  the  other  elements  consists  of  inosculating 
pits,  but  they  seldom  assume  the  form  of  ridges  or  grooves.  The  bones  of  the 
premaxillary  region  of  the  cranium  are  lacking.  The  first  element  which  can  be 
detected  is  the  prefrontal,  which  occupies  a  position  in  front  of  the  orbit.  There 
seems  to  be  space  for  a  lacrimal,  but  its  outline  is  not  distinct.  The  frontal  can  be 
readily  separated  and  is  seen  to  be  an  elongate  element  occupying  the  median  region 


THE    MICROSAURIAN   FAMILY   TUDITANID.«. 


87 


of  the  skull  between  the  orbits.  The  parietal  is  apparently  the  largest  element  of  the 
cranial  roof  and  the  pineal  foramen  is  located  in  the  anterior  fourth  of  the  median 
suture  separating  the  parietal  elements.  The  form  of  the  postparietal  and  the  tabu- 
lare  can  not  be  determined,  as  the  greater  part  of  this  region  is  lacking.  The  squa- 
mosal seems  to  be  located  well  forward  and  is  rather  small,  but  has  the  usual  rela- 
tion of  this  element.  Only  fragments  of  the  other  ele- 
ments remain  and  nothing  can  be  said  of  their  form. 
The  mandibles  of  both  sides  are  represented  by  depres- 
sions, and  they  are  ornamented  with  longitudinal  grooves 
and  ridges.  The  teeth  are  not  preserved,  but  there  are 
evidences  of  the  maxillary  teeth.  These  are  minute  and 
sharply  conical.  Just  posterior  to  the  skull  there  is  pre- 
served the  impression  of  a  short,  round  rod  which  is 
not  definitely  determined.  It  may  be  an  element  of  the 
hyoid  apparatus,  although  it  is  rather  stout  for  such.  It 
does  not  have  the  relations  indicated  by  Cope  in  his 
figure  (123,  pi.  xxxiv,  fig.  1). 

There  are  three  elements  of  the  pectoral  girdle  pre- 
served. These  undoubtedly  represent  the  interclavicle 
and  the  clavicles.  The  interclavicle  is  rhomboid  in  shape 
and  is  attenuated  posteriorly.  The  attenuation  is  ab- 
ruptly truncate  posteriorly  and  it  is  thus  of  quite  a  dif- 
ferent character  from  the  acutely  pointed  interclavicle 
of  T.  minimus.  The  clavicle  has  a  somewhat  semicir- 
cular form,  but  is  not  attenuated  at  either  end.  It  seems 
to  be  uniformly  broad. 

The  forearm  of  the  right  side  is  preserved  in  part. 
The  humerus  is  seen  to  be  a  heavy,  somewhat  expanded 
element  lying  displaced  with  relation  to  the  pectoral 
girdle.  It  is  greatly  expanded  at  the  ends.  The  ulna 
presents  characters  similar  to  the  humerus  and  only  dif- 
fers from  it  in  being  shorter  and  less  stout.  The  radius 
is  not  preserved.  The  carpus  is  unossified  and  its  posi- 
tion is  occupied  by  a  blank  space.  The  digits  are  repre- 
sented by  4  metacarpals,  and  this  may  have  constituted 
the  entire  number  of  the  fingers.  The  phalangeal  bones 
preserved  are  a  little  scattered.  They  are  elongate  with 
expanded  ends. 

Evidences  of  23  consecutive  osseous  vertebrae  are 
preserved.  Their  character  can  not  be  determined,  al- 
though Cope  (123)  describes  them  as  amphiccelous.  This  may  be  inferred  to  be  the 
case,  but  I  am  unable  to  verify  his  observation.  In  form  the  vertebras  are  subquad- 
rate.  The  neural  spines  are  not  evident.  The  osseous  ribs  articulate,  apparently, 
between  the  bodies  of  the  vertebrae.  Cope  figured  them  as  intercentral.  There  are  22 


Fig.  19. — Drawing  of  skull  and  skele- 
tal elements  of  Tuditanus  punctu- 
latus  Cope  from  the  Coal  Meas- 
ures of  Linton,  Ohio.  X  1.5.  fr, 
frontal;  ic,  interclavicle;  d,  clav- 
icle; h,  humerus;  ph,  phalanges; 
par,  parietal;  pp,  postparietal; 
lab,  tabulare,  supratemporal  and 
squamosal;  u,  ulna;  r,  radius. 


88  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

or  23  pairs  preserved.    They  are  single-headed  and  the  extremities  are  attenuated. 
No  traces  of  ventral  scutella  are  present. 

The  entire  length  of  the  animal  probably  did  not  exceed  5  or  6  inches.  Its 
form  was  quite  lizard -like  and  it  was  probably  of  an  ambulatory  type,  though  it 
may  have  spent  a  part  of  its  time  in  the  old  lagoon  in  which  its  remains  were  finally 
buried.  No  traces  of  external  gills  have  been  detected  in  this  or  any  other  Linton 
species. 

Measurements  of  the  Type  of  Tuditanus  punctulatus  Cope. 

ram.  mm. 

Length  of  entire  specimen 94  Width  of  humerus  at  proximal  end 2.5 

Median  length  of  skull 22  Length  of  ulna 6 

Width  of  skull  at  occiput 19  Width  of  ulna  at  proximal  end 2 

Depth  of  mandibular  ramus 4  Length  of  phalanx  (metacarpal?) 2 

Length  of  the  23  vertebra? 61  Expanse  of  longest  ribs 16 

Length  of  interclavicle 6.5  Length  of  rib 10.5 

Width  of  interclavicle 4.5  Width  of  rib 5 

Width  of  the  three  pectoral  plates 10  Length  of  a  vertebra 2.5 

Length  of  humerus 8 


Tuditanus  brevirostris  Cope. 

Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  272,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  393,  pi.  xxvi,  figs.  3,  4,  1875. 

Moodie,  Bull.  Amer.  Museum  Natl.  Hist.,  xxvi,  art.  xxv,  pi.  lxiv,  fig.  4,  1909. 

Type:  Specimen  No.  8609  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  species  was  associated  by  Cope  with  the  type  T.  punctulatus  in  the  descrip- 
tion of  the  genus.  Represented  by  a  portion  of  the  skeleton  of  one  individual,  the 
skull  is  preserved  on  one  block,  with  a  considerable  part  of  the  anterior  ribs,  pec- 
toral girdle,  and  vertebral  column,  although  this  last  is  not  clearly  represented,  but 
as  in  so  many  of  the  coal  specimens  the  bones  are  covered  with  a  thin  layer  of  car- 
bonaceous matter  which  makes  it  impossible  to  definitely  determine  the  form. 

The  cranium  is  large  in  proportion  to  the  size  of  the  body.  The  skull  is  in  the 
form  of  a  wide  oval  and  is  wider  than  it  is  long.  The  elements  of  the  skull  were  orna- 
mented with  a  coarse  sculpturing  which  partakes  of  the  nature  of  incomplete  radia- 
tions on  the  squamosal  region.  The  different  elements  of  the  cranium  can  not  be 
distinguished,  although  I  think  the  outlines  of  the  parietal  are  indicated.  The 
position  of  the  nostrils  is  well  forward  and  they  are  slightly  elongate  transversely. 
The  pineal  foramen  can  not  be  determined.  The  orbits  are  oval  in  shape  and  their 
width  is  about  equal  to  two-thirds  of  their  length.  The  interorbital  space  is  greater 
than  the  length  of  the  orbit.  Cope's  figure  of  this  specimen  is  not  accurate,  since 
he  has  the  orbits  drawn  too  far  to  the  side.  They  are  located  near  the  central  line 
of  the  skull  and  resemble  in  some  respect  those  of  the  preceding  species.  Cope  has 
described  teeth  in  the  maxillary  region,  but  I  am  unable  to  detect  them.  There  are 
portions  of  two  pectoral  elements  which  may  represent  a  clavicle  and  a  portion  of 
the  interclavicle. 

The  clavicle  has  much  the  same  shape  and  practically  the  same  ornamentations 
as  has  the  clavicle  in  Tuditanus  minimus.  The  clavicle  preserved  shows  a  somewhat 
triangular  shape  and  is  slightly  acuminate  at  the  anterior  end,  as  preserved,  and 
obtuse  at  the  posterior  end.    The  nature  of  the  interclavicle  can  not  be  determined. 


THE    MICROSAURIAN   FAMILY   TUDITANID^.  89 

The  vertebral  column  is  represented  by  a  line  which  Cope  suggests  (123)  may 
be  the  chorda  dorsalis  (notochord).  Osseous  vertebrae  were  probably  present,  but 
their  nature  is  obscured  by  the  carbonaceous  matter  covering  them. 

The  ribs  as  preserved  are  long  and  curved.  They  are  slender  and  attenuated  at 
the  distal  ends.  They  were  probably  single-headed,  but  whether  their  articulation 
was  intercentral  or  not  can  not  be  determined. 

The  other  specimens  which  are  referred  to  this  species  show  nothing  of  impor- 
tance in  the  way  of  structure.  They  consist  for  the  most  part  of  fragments  which 
may  or  may  not  represent  the  species. 

The  species  differs  from  the  type  of  the  genus  (T.  punctulatus)  in  the  possession 
of  a  broadly  rounded  muzzle.  This  character  will  also  separate  it  from  other  spe- 
cies of  the  genus.  The  sculpturing  of  the  bones  of  the  cranium  is  coarser  in  the  pres- 
ent species  than  in  the  type.  The  form  of  the  clavicle  is  different  in  the  two  species. 
The  above-described  species  seems  to  be  more  closely  allied  to  the  form  described 
as  Tuditanus  walcotti  than  to  other  species  of  the  genus.  I  have  been  unable  to 
detect  the  presence  of  limbs,  although  Cope  says  they  are  present. 

Measurements  of  the  Type  of  Tuditanus  brevirostris  Cope. 

mm.  mm. 

Median  length  of  skull 15         Interorbital  space 4.5 

Width  of  skull  at  posterior  border 18         Length  of  clavicle 7 

Width  of  skull  across  orbits 1 1 .5      Width  of  clavicle 3 

Length  of  orbit 3         Length  of  longest  rib  preserved 9 

Width  of  orbit 2         Length  of  entire  specimen 54 

The  material  consists  of  the  type  specimen  with  counterpart  and  two  fragments 
which  probably  are  to  be  associated  with  this  species.  Collected  by  Doctor  J.  S. 
Newberry. 

Tuditanus  longipes  Cope. 

Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  210,  1874  (Sauropleura). 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  II,  398,  pi.  xxvi,  fig.  2,  1875. 

Type:  Specimen  No.  1099  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  cranium  of  this  species  is  quite  unknown.  The  only  genus  with  which  the 
specimen  can  be  compared  in  the  structure  of  the  skeleton  is  Tuditanus.  From  the 
other  species  of  the  genus  the  present  form  differs  in  the  presence  of  ventral  chevron 
rods  and  the  elongate  character  of  the  limbs,  as  well  as  in  the  possession  of  large 
iliac  bones,  which,  in  the  only  other  species  in  which  the  ilia  are  known,  are  small 
and  slender.  It  seems  best  to  locate  the  species  in  this  genus  for  the  present, 
although  it  may  eventually  have  to  be  removed  to  another  group.  Very  little  is 
known  of  the  main  portion  of  the  skeleton  of  the  species  of  Tuditanus,  other  than  in 
T.  longipes,  so  an  exact  comparison  is  impossible.  From  all  the  species  of  Tuditanus 
thus  far  known  the  present  species  differs  in  the  elongate  character  of  the  limbs  and 
in  the  presence  of  ventral  scutellation.  There  are  three  other  species  of  Tuditanus 
in  which  the  limbs  are  known.  These  are:  T.  walcotti  Moodie,  T.  minimus  Moodie, 
and  T.  punctulatus  Cope.  In  these  three  species  the  limbs  are  short  and  weakly 
developed.  From  the  otherspecies  of  Tuditanus  the  present  species  maybe  separated 
by  its  size  principally,  since  nothing  of  the  bodies  of  the  other  species  is  known. 


90  THE    COAL   MEASURES   AMPHIBIA    OF   NORTH    AMERICA. 

The  body  of  the  present  species  is  elongate  and  slender,  with  a  long  neck  and  prob- 
ably a  long  tail.  Ribs,  as  preserved,  are  19  to  22,  though  there  may  possibly  have 
been  more.  They  are  moderately  curved  backwards,  have  intercentral  articulation, 
are  attenuated  at  the  distal  extremity,  and  are  single-headed.  The  anterior  ribs 
are  stouter,  with  a  widened  upper  portion  and  attenuated  distal  part.  The  posterior 
ribs  are  more  slender. 

There  are  evidences  of  28  vertebrae  present.  All  regions  of  the  vertebral  column 
are  present  and  the  dorsal  region  is  preserved  entire.  The  cervical  series  is  repre- 
sented by  the  posterior  vertebrae  only.  These  are  very  indistinctly  preserved.  The 
dorsal  vertebrae  are  elongate  and  were  prob- 
ably amphiccelous,  although  this  has  not 
been  definitely  determined.  They  are  ex- 
panded at  each  end,  thus  ending  in  a  slightly 
raised  rim.  The  single-headed  ribs  articulate 
between  the  vertebrae.  The  exact  number 
of  the  dorsal  series  can  not  be  ascertained, 
although  this  may  have  been  25.  The  spines 
of  the  vertebrae  are  not  determinable,  since 
the  animal  is  preserved  on  its  back.  The 
caudal  vertebrae  are  represented  by  two 
patches  of  the  remains  of  what  was  once 
probably  the  entire  series.  Cope  ascribes  70 
mm.  to  the  tail,  but  I  do  not  find  that  much. 
The  specimen  may  have  been  mutilated  since 
he  studied  it.  The  caudals  are  slender  and, 
like  the  dorsals,  are  expanded  at  the  ex- 
tremities. 

The  scapular  arch  is  not  preserved,  but 
the  pelvic  arch  is  represented  by  the  two  iliac 
bones  in  good  state  of  preservation.  These 
are  short,  flat  bones  expanded  at  the  anterior 
extremity,  as  preserved.  They  lie  turned  a 
little  to  each  side  of  the  vertebral  column  and    fig.  2o.— Cope's   drawing  of  TudUanus  longipes, 

partially  Obscure  the  femora.    The  iliac  bones  from  the  Linton,  Ohio,  Coal  Measures.     XI. 

are  quite  characteristic  of  this  form,  since  similar-shaped  elements  have  not  been 
observed  in  any  of  the  other  Carboniferous  forms  from  the  same  deposit. 

The  greater  part  of  the  forelimb  is  preserved,  although  much  of  the  hand  is 
missing.  The  humerus  is  an  unusually  elongate  bone  and  lies  somewhat  across  the 
vertebral  column.  It  is  crushed  flat  and  the  ends  are  partly  destroyed.  It  shows 
evidences  of  expansion  at  the  ends,  although  not  a  great  deal.  It  is  much  longer  than 
the  radius  and  ulna,  which  are  of  about  equal  length.  The  ulna  is  larger  than  the 
radius  and  has  expanded  ends,  with  the  upper  end  more  expanded  than  the  lower 
and  both  ends  slightly  truncate.  The  radius  is  a  simple  rod  of  bone  and  is  but 
slightly  expanded.    The  carpus  was  evidently  cartilaginous,  since  there  is  no  evi- 


THE   MICROSAURIAN    FAMILY   TUDITANID/E.  91 

dence  of  osseous  material  in  its  place.  There  is  but  one  phalangeal  bone  preserved, 
and,  since  this  is  displaced  with  reference  to  the  ulna  and  radius,  its  position  can 
not  be  determined.  It  may  have  been  a  metacarpal.  It  is  short  and  expanded  at 
the  ends. 

The  hind  limbs  are  represented  by  the  two  femora  and  the  upper  portion  of  the 
tibia.  The  femur  is  almost  as  elongate  as  the  humerus  and  is  more  slender.  It  is 
not  so  much  expanded  as  the  humerus.  Its  ends  appear  to  have  been  cartilaginous 
and  do  not  represent  the  well-formed  articular  surfaces  preserved  in  the  T.  minimus. 
The  upper  part  of  the  tibia  is  preserved,  and  appears  to  have  been  truncate. 

If  this  species  belongs  with  Tuditanus  it  is  of  interest  in  that  the  ventral  chev- 
rons are  present.    The  species  is  particularly  characterized  by  the  elongate  limbs. 

Measurements  of  the  Type  of  Tuditanus  longipes  Cope. 

mm.  mm. 

Length  of  vertebral  column  between  pelvis  and  end  Length  of  radius  and  ulna 12 

of  humerus 70      Length  of  a  vertebra 3 

Length  of  vertebral  column  anterior  to  humerus 18       Length  of  ilium 7 

Length  of  caudals  present 42      Length  of  femur,  estimated 18 

Length  of  humerus 19  7  chevrons  in  4  mm. 

Width  of  humerus 2 

Tuditanus  minimus  Moodie. 

Moodie,  Jour.  Geol.,  xvn,  No.  [,  p.  56,  fig.  10,  1909. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  23,  pi.  8,  fig.  2,  1909. 

Type:  Specimen  No.  4555,  U.  S.  National  Museum. 

Horizon  and  locality :  Cannelton  slates  of  Pennsylvania  (Upper  Freeport) . 

The  species  is  represented  by  a  nearly  complete  skeleton  preserved  on  a  slab  of 
slate  from  the  Cannelton  shales  of  Pennsylvania.  The  obverse  slab  has  been  lost, 
which  is  very  unfortunate,  since  there  is  no  doubt  that  the  entire  skeleton  was  origi- 
nally present.  The  species  is  placed  in  the  genus  Tuditanus  on  account  of  the  close 
resemblance  to  the  type  form  T.  punctulatus  Cope,  although  it  is  much  smaller  than 
that  species. 

The  type  specimen  of  the  species  did  not  attain  a  length  of  more  than  3.5  inches. 
Its  form  is  very  lizard-like,  but  its  structure  is  typically  amphibian.  The  form  of 
the  skull  is  especially  similar  to  that  of  the  type  species  T.  punctulatus,  which  it 
resembles  in  the  narrow  posterior  truncation  of  the  skull,  as  well  as  in  the  anterior 
position  of  the  orbits. 

The  skull  is  in  the  form  of  a  narrow  oval,  sharply  narrowed  posteriorly  and  trun- 
cate. The  orbits  are  located  well  forward  and  their  posterior  border  lies  in  front  of 
the  line  dividing  the  skull  transversely  into  equal  parts.  The  interorbital  space  is 
greater  than  the  diameter  of  the  orbit.  Impressions  of  teeth  are  preserved  on  the 
premaxilla?  and  maxillae;  there  are  8  of  them  in  a  distance  of  3  mm.  The  teeth 
appear  to  be  mere  blunt  denticles  and  were  possibly  pleurodont. 

The  elements  of  the  cranium  are  very  poorly  preserved.  It  has  been  impossible 
to  determine  all  of  the  sutures.  The  bones  of  the  premaxillary  region  have  been 
destroyed,  but  the  arrangement  of  them  was  probably  not  far  different  from  that 
which  obtains  in  other  members  of  the  genus.    The  posterior  boundaries  of  the  nasals 


92  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

are  preserved  and  prove  this  element  to  have  had  an  obtuse  posterior  border.  The 
sutures  bounding  the  frontals  are  clear  and  show  that  they  were  small  and  that  they 
formed  a  part  of  the  inner  boundary  of  the  orbits.  The  parietal  is  recognized  as  a 
large  element,  apparently  the  largest  in  the  skull.  Together  the  parietals  form  a 
wide  oval  inclosing,  on  the  median  suture,  the  circular  pineal  foramen.  The  parie- 
tals are  sculptured  with  coarse  radiating  grooves  and  ridges,  much  after  the  manner 
of  Erpetosaurus  radiatus  Cope.  The  pittings  present  on  that  form  are,  however, 
absent  in  T.  minimus.  The  sutures  bounding  the  postparietal  are  tolerably  well 
defined  and  these  show  that  element  to  have  been  rather  large  and  quadrate,  with 
the  usual  relations.  The  tabulare  is  distinct,  triangular,  and  small.  It  is  produced 
into  an  angle  on  the  posterior  border  strongly  recalling  a  similar  condition  in  T. 
punctulatus.  The  boundaries  of  the  prefrontals  and  the  upper  borders  of  the  max- 
illae are  not  clearly  ascertained.  The  lacrimal  has  not  been  detected.  The  post- 
frontal  and  postorbital  form  the  posterior  boundary  of  the  orbit,  although  all  of  the 
limits  of  the  latter  element  have  not  been  definitely  determined.  The  position  of 
the  supratemporal  is  well  assured,  although  its  entire  boundaries  are  not  determined. 
It  has  the  usual  relations  and  joins  the  parietal  broadly.  The  jugal  is  broad  and 
widens  posteriorly  to  join  the  squamosal,  which,  as  usual,  forms  the  quadrate  angle 
of  the  skull.  The  sutures  bounding  the  quadratojugal  and  the  posterior  end  of  the 
maxilla  are  not  determined. 

There  are  but  two  fragmentary  vertebras  preserved  and  an  estimate  based  on  the 
length  of  these  remains  gives  about  30  presacral  vertebrae.  The  structure  of  the 
vertebrae  preserved  can  not  be  ascertained,  but  the  neural  spines  appear  to  have 
been  low  and  stout. 

There  are  six  elements  of  the  pectoral  girdle  preserved.  These  are:  the  six 
clavicles,  the  interclavicle,  the  coracoid  of  one  side,  and  the  two  scapulas.  The 
interclavicle  is  rhomboid  in  form  and  acuminate  posteriorly.  It  is  sculptured 
with  radiating  grooves  and  ridges.  It  is  quite  different  from  the  same  element 
in  T.  punctulatus,  in  that  the  base  is  acuminate,  not  truncate.  The  clavicle 
presents  much  the  same  shape  as  does  that  element  in  Erpetosaurus  tabulatus.  It  is 
ornamented  by  a  sculpturing  of  radiating  lines  which  take  their  origin  from  the 
lower  external  angle  as  the  bone  lies  in  the  matrix.  The  clavicle  is  somewhat  trian- 
gular in  shape  and  lies  close  to  the  skull,  but  this  close  approximation  of  the  pecto- 
ral elements  to  the  cranium  is  due  probably  to  post-mortem  shifting,  since  the 
scapulae  are  shifted  backward.  There  can  be  little  doubt,  however,  that  the  pectoral 
arch  was  close  to  the  cranium.  There  is  an  oval  fragment  preserved  on  the  left  of 
the  specimen  which  I  take  to  be  a  portion  of  the  coracoid.  The  scapula  is  preserved 
entire  on  the  left  side  and  is  represented  by  fragments  on  the  right  side.  It  is 
almost  semicircular  in  form  and  narrows  externally  until  it  is  somewhat  fan-shaped. 
There  appears  to  be  an  ornamentation  of  lines  on  the  surface  of  the  bone.  These 
lines  follow  the  contour  of  the  anterior  border. 

The  arm  is  preserved  nearly  complete  on  the  left  side,  and  the  right  side  shows 
the  humerus  and  the  forearm.     The  humeri  are  unusual  in  having  well-developed 


THE    MICROSAURIAN    FAMILY   TUDITANID^E.  93 

articular  ends  as  though  the  endochondral  tissue  was  well  developed.  The  hume- 
rus is  expanded  at  the  ends  and  it  is  larger  at  the  upper  than  at  the  lower  end. 
The  ulna  is  expanded  at  the  proximal  extremity,  but  is  more  attenuated  at  the  distal 
portion.  It  is  shorter  than  the  humerus  by  about  one-third  of  its  own  length.  The 
radius  is  a  mere  slender  rod  of  bone  and  presents  well-developed  articular  ends.  It 
is  slightly  shorter  than  the  ulna.  The  carpus  is  unossified  and  its  position  is  repre- 
sented by  a  blank  space.  There  are  phalanges  of  4  digits  preserved  and  they  are  4 
in  number.  The  phalangeal  elements,  like  the  other  bones  of  the  extremity,  have 
the  articular  surfaces  prominent,  with  the  terminal  phalanx  claw-like. 

There  are  no  ribs  nor  traces  of  them  preserved,  and  a  conjecture  as  to  their 
character  can  not  be  hazarded,  since  they  are  known  in  but  two  other  species,  in 
which  they  are  slender  and  curved.  There  is  no  evidence  of  a  ventral  scutellation, 
and  so  far  as  is  at  present  known  this  structure  is  absent  from  all  of  the  species  of 
the  genus,  or  at  least  it  is  weakly  developed. 

The  ilium  is  all  that  is  preserved  of  the  pelvis.  The  bone  itself  has  disappeared 
and  has  left  a  depression  which  shows  this  element  to  have  been  an  elongate  rod  very 
similar  to  that  described  for  Micrerpeton.  The  sacral  vertebra  seems  to  be  indicated 
by  a  depression  between  the  iliac  depressions. 

One  hind  limb  is  preserved  nearly  entire  and  the  greater  part  of  the  other  is  also 
preserved,  although  some  of  the  phalanges  are  disturbed.  The  femur  is  slender  and 
more  elongate  than  the  humerus.  It  has  well-formed,  rounded,  articular  ends.  The 
tibia  presents  unusual  characters  in  that  its  ends  are  truncate,  as  though  the  car- 
tilage composing  its  articular  surfaces  was  not  so  highly  calcified  as  in  the  other 
limb  bones.  It  is  somewhat  expanded  at  the  ends  and  is  throughout  its  length 
broader  than  the  femur.  The  fibula,  like  the  tibia,  is  a  slender  rod  of  bone,  although 
it  is  somewhat  shorter  than  the  tibia.  The  tarsus  is  unossified  and  its  position  is 
occupied  by  a  blank  space.  Portions  of  both  feet  are  preserved,  but  only  one  digit 
in  the  right  foot  is  complete.  The  metatarsals  are  elongate  and  slightly  expanded 
at  the  ends.  There  are  4  phalanges  present  in  the  complete  digit,  which  possibly 
represents  the  fourth.  The  first  digit  is  wanting,  with  the  only  terminal  phalanx 
preserved  claw-like. 

Measurements  of  the  Type  of  Tuditanus  minimus  Moodie. 

mm.  mm. 

Median  length  of  skul! 15  Width  of  scapula,  maximum 2.5 

Width  of  skull  at  posterior  border 16  Length  of  coracoid  (?) 2 

Length  of  orbit 3.5  Length  of  humerus 4 

Width  of  orbit 2  Length  of  radius  and  ulna 3 

Interorbital  width 2.5  Length  of  metacarpal I 

Length  of  clavicle 6  Length  of  ilium 2.5 

Width  of  clavicle,  maximum 3.5  Length  of  femur 4.5 

Length  of  interclavicle,  estimated 5  Length  of  tibia  and  fibula 3 

Width  of  interclavicle 3.5  Length  of  foot ; 3.5 

Length  of  scapula 3.5  Length  of  metatarsal 75 

Tuditanus  walcotti  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  xxxvn,  p.  16,  pi.  6,  figs.  1,  2;  pi.  7;  1909. 

Type:  Specimen  No.  4474,  U.  S.  National  Museum. 
Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 


94 


THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


A  small  species  of  Microsauria  is  preserved  as  a  smooth  impression  on  a  block  of 
soft  coal  from  Linton,  Ohio.  Nearly  the  entire  form  of  the  body  is  discernible.  The 
specimen  is  especially  interesting  and  valuable  as  exhibiting  for  the  first  time  among 
the  Linton  forms  the  shape  of  the  body  of  the  small  microsaurians  of  the  Tuditanus 
type.  It  differs  so  markedly  in  the  form  of  the  skull  from 
other  species  of  the  genus  that  it  is  regarded  as  a  distinct 
form,  and  the  name  Tuditanus  walcotti  was  proposed  for 
it  as  an  expression  of  the  writer's  indebtedness  to  the 
Secretary  of  the  Smithsonian  Institution  for  the  use  of 
the  material  among  which  the  present  form  was  included. 

The  specimen  includes,  besides  the  body  impression, 
the  complete  skull,  a  right  clavicle,  with  portions  of  the 
left,  a  left  humerus,  12  cervical  and  dorsal  vertebrae,  10 
pairs  of  ribs  somewhat  disturbed  as  to  position,  and  a 
portion  of  the  mandible.  There  are  no  traces  of  ventral 
scutellae  nor  body  scales  in  the  smooth  impression  of  the 
carbonized  skin.  One  would  expect  to  find  impressions 
of  the  ventral  scutse  in  this  specimen  if  they  were  present. 
Cope  remarked  on  the  apparent  absence  of  scutellae  from 
members  of  the  genus  Tuditanus  as  they  were  known  to 
him,  and  no  contrary  evidence  has  since  been  brought 
to  light.  Until  such  evidence  is  forthcoming  the  absence 
of  scutes  will  be  taken  as  one  of  the  generic  characters 
of  the  genus  Tuditanus.  Under  a  magnification  of  50 
diameters  the  carbonized  skin  shows  as  folds  and  wrin- 
kles, like  muscle  fibers,  in  some  places ;  in  others  no  traces 
of  the  muscular  structure  can  be  detected.  The  wrinkles 
may  be  impressions  of  the  internal  musculature  of  the 
body -wall  of  the  abdomen.  It  is  especially  well  preserved 
in  the  pelvic  and  pygal  regions.  Sections  of  the  coal 
were  made,  but  nothing  definite  could  be  determined  as 
to  the  character  of  the  impressions,  as  they  were  too 
poorly  preserved  and  the  coal  was  too  soft  to  bear  much 
handling. 

The  specimen  is  preserved  on  the  belly,  with  the 
dorsum  of  the  skull  uppermost.  It  has  been  practically 
impossible  to  determine  the  arrangement  of  any  of  the 
cranial  elements  except  the  frontals,  parietals,  and 
postparietals,  which  have  the  relations  indicated  in 
figure  21,  A.  A  median  suture  is  clearly  evident,  with  the  pineal  foramen  well  back 
in  this  suture.  The  bones  of  the  skull  are  marked  with  faint  radiating  lines.  It 
is  in  the  form  of  the  skull  and  the  position  of  the  orbits  that  the  specific  char- 
acters are  found,  as  follows:  the  backward  position  of  the  eyes  and  the  oval, 
pointed  shape  of  the  skull.     The  species  is  closely  related  to  Tuditanus  minimus 


Fig.  21. —  A.  Outline  drawing  of  type 
of  Tuditanus  walcotti  Moodie, 
from  the  Coal  Measures  of 
Linton,  Ohio,  showing  impres- 
sion of  body  and  muscle  at  M. 
X  25.  cl,  clavicle;  fr,  frontal;/, 
femur;  h,  humerus;  nos,  nostril; 
or,  orbit;  par,  parietal;  rb,  rib; 
pp,  postparietal ;  v,  vertebra;  pfo, 
pineal  foramen. 

B.  Left  leg  of  second  specimen  of 
Tuditanus  walcotti.     X  3. 


THE    MICROSAURIAN    FAMILY    TUDITANID/E.  95 

Moodie,  from  the  Cannelton  slates  of  Pennsylvania,  and  serves  further  to  connect 
the  forms  from  the  Ohio  and  Pennsylvania  localities.  It  differs  from  the  last-named 
species  in  the  position  and  form  of  the  orbits,  these  structures  being  more  oval  in  the 
present  form  and  placed  further  back.  The  shape  of  the  skull  differs  also  in  the 
almost  entire  absence  of  the  posterior  table.  The  median  points  of  the  orbits  occupy 
the  line  which  bisects  the  skull,  and  the  interorbital  width  is  less  than  the  width  of  the 
orbit.     The  mandible  is  heavy  and  appears  to  have  borne  sharp,  pleurodont  teeth. 

The  vertebral  column  is  represented  by  little  more  than  a  mold  of  the  form  of 
the  vertebrae,  so  that  little  can  be  said  of  its  character.  The  individual  vertebrae  are 
short  and  hour-glass-shaped.  The  ribs  are  borne  intercentrally,  as  in  all  the  micro- 
saurians  which  have  been  studied  from  the  Linton  deposits.  The  ribs  are  rather 
long  and  somewhat  heavy,  slightly  curved  and  expanded  at  the  proximal  end,  as 
though  an  incipient  bicipital  condition  were  present. 

The  right  clavicle,  which  is  preserved  as  an  impression,  is  entire.  Its  impres- 
sion shows  this  element  to  have  been  ornamented  on  its  ventral  surface  with  radi- 
ating grooves  and  ridges  which  started  at  the  lower  angle  of  the  bone.  The  element 
is  distinctly  triangular,  which  is  characteristic  of  the  genus  Tuditanus,  so  far 
as  known.  The  fragment  of  the  left  clavicle  adds  nothing  to  our  knowledge  of 
the  element. 

The  left  humerus  recalls  in  a  striking  way  that  of  Tuditanus  longipes  Cope,  and 
it  was  once  entertained  as  a  possibility  that  the  present  form  might  be  a  member  of 
that  species,  since  the  skull  is  lacking  in  T.  longipes.  Sufficient  specific  differences 
were  found,  however,  in  the  ribs,  which,  in  T.  longipes,  are  very  long,  slightly  curved, 
and  delicate,  but  which,  in  the  present  form,  are  comparatively  heavy.  Other  char- 
acters sufficiently  diagnostic  are  found  in  the  form  assumed  by  the  vertebrae  in  the 
two  forms. 

Measurements  of  the  Type  of  Tuditanus  walcotti  Moodie. 

mm.  mm. 

Length  of  specimen 70  Length  of  vertebral  column,  as  preserved 50 

Length  of  skull 20  Length  of  a  vertebra 1.75 

Posterior  width  of  skull 14  Width  of  a  vertebra 50 

Width  of  skull,  anterior  to  orbits 10  Width  of  body  impression 15 

Length  of  orbit 4  Length  of  humerus 6 

Width  of  orbit 2  Median  width  of  humerus 50 

Interorbital  width 3  Width  at  end  of  humerus 2 

Length  of  clavicle 9  Length  of  rib 8 

Greatest  width  of  clavicle 4  Width  of  rib 25 

The  above-described  specimen  was  collected  by  Mr.  R.  D.  Lacoe,  of  Pittston, 
Pennsylvania,  from  Linton,  Ohio. 

A  second  individual  (No.  4481,  U.  S.  National  Museum)  of  this  species  is  indi- 
cated by  a  rather  poorly  preserved  specimen  on  a  slab  of  soft  coal  from  the  Linton 
mines.  The  following  portions  of  the  animal  have  been  detected  and  will  be  dis- 
cussed: partial  impression  of  the  skull,  with  a  fragment  of  a  minute  jaw,  in  which 
are  minute  teeth ;  right  clavicle ;  part  of  the  impression  of  the  body ;  nearly  entire 
left  hind  limb ;  impressions  of  about  a  dozen  vertebrae,  very  indistinct. 

The  impression  of  the  skull  is  distinct  only  in  a  favorable  light,  and  even  then  the 
boundaries  of  the  cranium  are  a  little  uncertain.    For  this  reason  no  representation 


g6  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

of  the  form  will  be  attempted.  The  sculpturing  on  the  parietals  is,  however,  dis- 
tinct enough  to  show  relationship  with  the  previously  described  specimen,  and  the 
form  of  the  body  impression,  the  absence  of  abdominal  scutes,  the  shape  of  the  clavi- 
cle and  its  sculpture,  and  the  proportions  of  the  hind  limbs  all  agree  with  the  charac- 
ters which  have  been  assigned  to  the  genus  Tuditanus.  The  fragment  of  the  jaw  is 
interesting  as  giving  the  first  information  as  to  the  character  of  the  mandible  in  the 
genus  Tuditanus.  It  is  very  slender  and  of  uniform  width  as  far  as  preserved.  The 
teeth  are  short,  blunt  cones,  apparently  pleurodont. 

The  clavicle  is  of  the  typical  Tuditanus  form,  with  the  sculpturing  lines  radiating 
out  from  the  angle.  The  impression  of  the  body  adds  nothing  to  that  already 
described  for  the  type  specimen.  The  nearly  entire  hind  limb  is  of  great  interest  as 
adding  another  example  of  the  phalangeal  formula.  The  foot  is  almost  perfectly 
preserved,  and  the  formula  was  probably  2-2-3-3-2.  The  endochondrium  of  the 
limb  bones  is  not  highly  developed.  About  a  dozen  vertebrae  are  represented  by 
molds  in  the  soft  coal,  but  nothing  of  their  structure  can  be  determined. 

The  sharp,  reptile-like  claws  in  which  the  toes  end  (fig.  2 1 ,  B)  recall  those  of  Eosau- 
ravus  and  of  Tuditanus  minimus  Moodie.  It  is  another  link  in  the  chain  of  the 
suggested  relationship  between  the  microsaurians  and  the  early  reptiles. 

Measurements  of  the  Second  Specimen  of  Tuditanus  walcotti  Moodie. 

mm.  mm. 

Length  of  entire  body  impression 75  Length  of  clavicle 8 

Width  across  belly,  maximum 16  Width  of  clavicle,  maximum 4 

Length  of  skull ?I7  Length  of  hind  limb 22 

Posterior  width  of  skull ?I4  Length  of  femur 8 

Length  of  fragment-of  jaw 4  Length  of  tibia  (?) 6 

Width  of  fragment  of  jaw 1.5  Length  of  metatarsal 2 

Length  of  tooth  in  jaw .25  Length  of  first  digit 6 

Genus  ERPETOSAURUS  Moodie,  1909. 

Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  p.  348,  fig.  1,  1909. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  21,  1909. 

Type:  Erpetosaurus  radiatus  Cope. 

Skull  stout,  elements  sculptured  with  radiating  grooves,  ridges,  and  pits;  orbits 
large  and  usually  placed  far  forward ;  occiput  sometimes  with  posterior  table ;  skull 
more  or  less  rounded;  lateral-line  canals  consisting  of  supraorbital,  suborbital,  jugal, 
and  temporal  canals,  the  last  two  uniting  to  form  a  circular  canal  in  one  species; 
clavicle  triangular,  sculptured  like  the  skull. 

Our  knowledge  of  the  genus  is  confined  to  the  skull.  The  genus  was  established 
for  certain  members  of  the  genus  Tuditanus  and  other  forms  which  have  been  re- 
cently described.  The  species  of  the  genus  are:  E.  radiatus  Cope  type,  E.  tabulatus 
Cope,  E.  tuberculatus  Moodie,  E.  obtusus  Cope,  E.  minutus  Moodie,  E.  acutirostris 
Moodie,  E.  sculptilis  Moodie.  All  of  the  species  are  from  the  Linton,  Ohio,  Coal 
Measures,  with  the  exception  of  E.  sculptilis  and  E.  minutus,  which  are  from  the 
Cannelton,  Pennsylvania,  slates. 

The  position  of  the  genus  as  to  family  is  a  little  uncertain,  since  family  charac- 
ters are  not  yet  well  understood  among  the  Carboniferous  forms  on  account  of  the 
lack  of  information  as  to  the  structure  of  the  animals.    If  we  take  the  absence  of 


THE   MICROSAURIAN   FAMILY   TUDITANID^E.  97 

the  ventral  scutellae  as  a  family  character,  the  genus  will  be  in  the  family  Tuditan- 
idae,  but  the  evidence  on  this  point  is  negative.  For  the  present  we  may  place  the 
genus  only  provisionally  in  the  family  Tuditanidae.  The  arrangement  will  undoubt- 
edly require  revision  later. 

Erpetosaurus  radiatus  Cope,  1874. 
Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  273,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  pp.  394-395,  pi.  xxvii,  fig.  1;  pi.  xxxiv,  fig.  3;  text-fig.  10,  1875. 
Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  p.  348,  pi.  lxii,  fig.  1,  1909. 

Type :  Specimen  No.  8600  G,  American  Museum  of  Natural  History. 
Locality  and  horizon:  Linton,  Ohio,  Coal  Measures.     (Plate  25,  fig.  1.) 
Cope  originally  described  this  species  from  a  portion  of  a  skull.    He  (123)  char- 
acterized the  form  as  follows: 

"The  marked  character  of  this  form  is  seen  in  the  very  anterior  position  of  the  orbits 
and  the  contraction  of  the  muzzle.  The  orbits  are  large  and  are  separated  by  a  little 
more  than  their  own  diameter;  their  posterior  border  is  in  front  of  a  line  measuring  the 
anterior  third  of  the  length  to  the  supraoccipital  crest,  and  nearly  to  the  line  marking  the 
fourth  of  the  length  to  the  quadrate  region.  The  posterior  outline  of  the  skull  is  deeply 
concave,  the  quadrate  angle  projecting  beyond  the  occipital  condyles." 

The  base  of  the  specimen  is  broken  and  there  is  no  place  for  the  occipital  condyles. 
Unless  the  specimen  has  been  mutilated  since  Cope  studied  it,  the  occipital  condyles 
are  not  present. 

The  restoration  of  the  skull  given  in  figure  22,  B  varies  but  little  from  that  given 
by  Cope  in  1875.    The  elements  are  practically  as  he  represented  them. 

The  premaxilla?  are  small  and  lie  in  the  usual  relations  to  the  other  elements. 
Minute  conical  teeth  are  present  as  impressions.  They  are  quite  similar  to  the  teeth 
found  in  other  Microsauria.  The  nasals  are  nearly  square  and  form  the  inner  boun- 
dary of  the  somewhat  oval  nostril,  which  is  represented  by  a  depression  in  the  coal. 
The  frontal  is  elongate.  It  is  about  twice  as  long  as  wide.  It  forms  a  portion  of  the 
inner  border  of  the  orbit,  the  remainder  being  made  up  by  the  prefrontals  and  the 
postfrontal.  The  parietals  are  the  largest  elements  of  the  skull,  but  they  do  not 
greatly  exceed  the  jugals.  Together  the  parietals  form  a  somewhat  obtuse  oval 
in  the  median  region  of  the  skull  and  they  contain  between  them,  in  their  posterior 
third,  the  small  circular  pineal  foramen.  The  postparietal  forms  the  posterior 
boundary  of  the  skull.  The  prefrontal  forms  the  anterior  border  of  the  orbit  and 
is  triangular  in  shape.  The  lacrimal  is  not  identified.  The  maxilla  is  an  elongate 
element  the  boundaries  of  which  are  uncertain,  though  probably  somewhat  as  given. 
The  postfrontal  and  the  postorbital  form  the  posterior  boundary  of  the  orbit, 
inclosing  between  them  the  anterior  projection  of  the  squamosal.  The  squamosal  is 
an  elongate  element  and  is  acuminate  at  each  end.  The  tabulare  is  a  large  element 
lying  lateral  to  the  postparietal.  The  jugal  is  a  very  elongate  element,  apparently 
acuminate  anteriorly.  The  quadratojugal  is  small  and  elongate.  The  supratemporal 
is  definitely  bounded  and  its  limits  are  as  indicated  (fig.  22  B),  being  a  large  element 
which  forms  the  quadrate  angle. 

There  are  two  other  specimens  of  this  species  in  the  collections  and  a  fragment 
of  a  fourth  which  it  is  difficult  to  make  out.    Cope  identified  and  figured  one  of  these 


98  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

as  E.  radiatus  (Geol.  Surv.  Ohio,  II,  pt.  II,  pi.  34,  fig.  3),  but  the  identification  is 
doubtful  and  the  figure  shows  structures  which  I  am  unable  to  identify  in  the  speci- 
men. Nothing  of  importance  is  to  be  learned  from  the  other  two  specimens,  except 
that  they  show  a  diversity  of  size.  They  consist  of  incomplete  skulls,  concerning 
which  Cope  (123)  remarks: 

"There  are  no  mucous  canals.  The  sculpture  consists  of  strong  ridges  radiating  and 
inosculating.  Radiation  is  more  uninterrupted  on  both  jugal,  supratemporal,  and  anterior 
part  of  the  tabulare;  on  the  first  they  originate  in  front  of  the  middle  exteriorly;  on  the 
supratemporal  near  the  anterior  part.  The  inosculation  is  honeycomb-like  on  the  parietal, 
postfrontal,  and  posterior  parts  of  the  tabulare." 

Measurement  of  the  Type  Specimen. 

mm.  mm. 

Length  of  the  skull  along  median  line  (estimated) ...   60      Length  of  orbit 9 

Length  from  muzzle  to  quadrate  angle 71       Width  of  orbit 8 

Width  at  posterior  border 69      Interorbital  width 8 

Width  at  orbits 40      Length  of  nostrils 2 

Measurements  of  Another  Specimen  of  the  Species. 
(No.  8598  G,  American  Museum  of  Natural  History.) 

mm.  mm. 

Median  length  of  skull 56      Length  of  orbit 7 

Length  to  quadrate  angle 61       Width  of  orbit 6 

Width  at  posterior  border 50      Interorbital  width 4.5 

The  specimens  of  this  species  were  collected  by  Dr.  J.  S.  Newberry. 

Erpetosaurus  obtusus  Cope,  1868. 

Cope,  Proc.  Phil.  Acad.  Nat.  Sci.,  1868,  p.  213. 

Cope,  Trans.  Amer.  Phil.  Soc,  xiv,  p.  12,  fig.  I,  1869. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  396,  fig.  11,  1875. 

Cope,  Proc.  Amer.  Phil.  Soc,  1885,  xxn,  p.  407  (Pal.  Bull.  40). 

Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  p.  350,  pi.  lx,  fig.  2. 

Type:  Specimen  No.  8601  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species,  Tuditanus  obtusus,  was  first  described  by  Professor  Cope  as  Den- 
drerpeton  obtusum,  but  he  subsequently  referred  the  species  to  the  genus  Tuditanus. 
It  was  removed  by  the  writer  to  the  genus  Erpetosaurus  in  1909.  The  species  is 
known  from  two  partially  preserved  crania.  The  skull  elements  seem  to  have  disap- 
peared and  left  only  the  impressions.  The  sutures  are,  for  the  most  part,  clearly 
represented,  but  the  skull  shows  no  sculpturing.  On  the  posterior  third  of  one 
cranium  there  is  a  small  space  which  seems  to  be  slightly  sculptured,  as  Cope  indi- 
cated in  his  drawing.  The  general  form  of  the  skull  is  that  of  a  broad  oval,  truncate 
posteriorly.  The  orbits  lie  in  the  anterior  third  of  the  skull,  and  the  pineal  foramen 
in  the  posterior  third.  Cope  compared  the  skull  to  that  of  Huxley's  Erpetocephalus, 
to  which  it  has  some  resemblance.  The  nostrils  are  elongate  and  are  situated  at  an 
obtuse  angle  with  relation  to  the  main  axis  of  the  skull. 

The  premaxilla  is  small  and  forms  the  inner  border  of  the  nostril.  There  seem 
to  be  impressions  of  small  teeth,  but  no  large  ones  are  evident.  The  nasals  are  sepa- 
rated by  a  zigzag  suture,  and  are  nearly  square.  They  have  the  usual  relations. 
The  frontals  form  a  portion  of  the  inner  boundary  of  the  orbits  and  unite  behind 


THE   MICROSAURIAN   FAMILY   TUDITANID/E. 


99 


with  the  parietals,  the  anterior  extensions  of  which  they  inclose.  The  parietals  are 
large  elements  and  together  form  a  broad  oval,  truncate  posteriorly.  They  inclose 
between  them  the  pineal  foramen  in  the  median  suture.  It  lies  in  the  posterior 
fourth  of  the  parietals.  The  postparietals  are  elongate  transversely,  and  have  the 
usual  relations.  The  prefrontal  is  somewhat  triangular  and  forms  the  anterior 
boundary  of  the  orbit.    The  lacrimal  has  not  been  detected.    The  maxilla  is  elon- 


FlG.    22. 

A.  (  hillinc  of  skull  and  cranial  elements  of  Erpetosaurus  minulus  Moodic,  from  the  Cannelton  slates  of  Pennsylvania. 

Original  in  U.  S.  National  Museum.  X  2.  fr,  frontal;  mx,  maxilla;  par,  parietal;  po,  postorbital;  pp,  post- 
parietal;  pr,  postfrontal;  sq,  squamosal;  spt,  supratemporal;  tab,  tabulare. 

B.  Outline  of  skull  and  cranial  elements  of  Erpetosaurus  radialus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio; 

partially  restored.  X  0.75.  Original  in  American  Museum  of  Natural  History,  fr,  frontal;./,  jugal;  mx,  maxilla; 
«,  nasal;  or,  orbit;  par,  parietal;  pf,  postfrontal;  po,  postorbital;  pr,  prefrontal;  pp,  postparietal ;  pmx,  premaxilla; 
qj,  quadra  to  jugal;  sq,  squamosal;   spt,  supratemporal;  tab,  tabulare. 

C.  Palate  of  Erpetosaurus  {tabulatus?) ,   from  the  Coal  Measures  of  Linton,  Ohio.     X  1.     American   Museum  of 

Natural  History,  ex,  exoccipital;  m,  mandible;  mx,  maxilla;  pal,  palatine;  pt,  pterygoid;  prv,  prevomer;  pd,  para- 
sphenoid  ;  *,  anterior  and  posterior  palatine  vacuities. 

D.  Outline  of  skull  and  cranial  elements  of  Erpetosaurus  acutiroslris  Moodie,  from  the  Coal  Measures  of  Linton, 

Ohio.   Original  in  American  Museum  of  Natural  History.     X  I.   ex,  exoccipital ;fr,  frontal;  j,  jugal;  mx,  maxilla; 
n,  nasal;  or,  orbit;  po,  postorbital;  par,  parietal;  pp,  postparietal;  pf,  postfrontal;  sq,  squamosal;  spt,  supra- 
temporal; tab,  tabulare. 
B,  ( Hitline  of  larger  part  of  skeleton  of  Odonterpeton  triangularis  Moodie,  from  the  Coal  Measures  of  Linton,  Ohio. 
Original  in  U.  S.  National  Museum.    X  4.    h,  humerus;  il,  ilium?;  v,  vertebrae,  all  of  which  are  not  represented. 

F.  Right  mandible  of  Erpetosaurus  tabulatus  Cope,  from  the  Linton,  Ohio,  Coal  Measures.    Original  in  American 

Museum  of  Natural  History.     X  2.    art,  articular;  cor,  coronoid;  d,  dentary;  ang,  angular;  sa,  surangular. 

G.  Skull  elements  and  lateral-line  canals  of  Erpetosaurus  tabulatus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio. 

X  1.5.  fr,  frontal;  j,  jugal;//,  jugal  lateral-line  canal;  mx,  maxilla;n,  nasal;  or, orbit;  par,  parietal;  po, postorbital ; 
pf,  lacrimal  or  prefrontal ;  so,  suborbital  lateral  -line  canal;  pp,  postparietal;  pmx,  premaxilla ;  qj,  quadratojugal ; 
spo,  supraorbital  lateral-line  canal;  sq,  squamosal;  spt,  supratemporal;  tm,  temporal  lateral-line  canal;  tab,  tabu- 
lare; so,  supraoccipital  cross-commissure  of  lateral-line  canal. 


IOO  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

gate,  and  with  the  quadratojugal  forms  the  exterior  border  of  the  cranium.  No 
teeth  are  observed  on  the  maxilla.  The  postfrontal  and  the  postorbital  form  the 
posterior  border  of  the  orbit,  and  between  them  inclose  the  anterior  extension  of  the 
squamosal.  The  supratemporal  is  pointed  anteriorly  and  has  the  usual  relations  of 
that  element,  joining  the  postorbital,  the  postfrontal,  the  parietal,  the  tabulare, 
and  the  squamosal.  The  tabulare  is  larger  than  the  postparietal  and  is  acuminate, 
the  point  being  inclosed  by  the  squamosal  and  the  supratemporal.  The  jugal 
widens  fan-shaped  posteriorly.  It  forms  a  portion  of  the  border  of  the  orbit. 
The  supratemporal,  as  usual,  forms  the  quadrate  angle  of  the  skull.  In  front  of  it 
lies  the  elongate  quadratojugal. 

Dendrerpeton  is  not  well  enough  known  for  an  exact  comparison  with  Erpeto- 
saurus  obtusus,  but  Cope  separated  the  latter  from  Dendrerpeton  on  the  position  of 
the  orbits  and  the  broadly  rounded  muzzle.  This  species  differs  from  the  other 
species  of  the  genus  in  the  form  of  the  cranium,  as  well  as  in  the  characters  which 
separate  it  from  Dendrerpeton. 

Measurements  of  the  Type  of  Erpetosaurus  obtusus  Cope. 

mm.  mm. 

Median  length  of  the  skull 48  Width  of  orbit 7 

Width  of  skull  at  posterior  border 56  Interorbital  space 15 

Width  across  orbits 37  Diameter  of  nostril 1.5 

Length  of  orbit 12.5  Diameter  of  pineal  foramen 1.75 

Two  other  specimens,  Nos.  8602  G  and  8608  G  of  the  American  Museum,  are 
associated  in  this  species. 

Erpetosaurus  tabulatus  Cope. 

Cope,  Proc.  Amer.  Phil.  Soc,  xvi,  p.  577  (Tudilanus  tabulatus). 

Moodie,  Jour.  Geol.,  17,  p.  52,  figs.  8,  9,  1909. 

Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  pp.  347,  351,  pi.  59,  fig.  2;  pi.  62,  fig.  2,  1909. 

Type:  Specimen  in  the  Zoological  Collection  of  Columbia  University. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.     (Plate  25,  fig.  2.) 

The  species  is  known  from  a  single  well-preserved  skull  and  its  obverse  in  the 
collection  of  Columbia  University  in  New  York  City.  I  am  indebted  to  Dr.  Bash- 
ford  Dean  for  the  privilege  of  studying  this  interesting  form.  It  is  from  the  Linton 
deposits  of  Ohio.  The  remains  include  a  nearly  complete  cranium  and  a  complete 
clavicle  of  the  right  side.  The  species  agrees  in  all  essential  respects  with  the  char- 
acters of  the  genus  Erpetosaurus,  presenting  a  broad,  flat  head  and  a  triangular 
clavicle. 

The  cranium  is  wider  than  long,  the  muzzle  broadly  rounded.  The  orbits  are 
wide  ovals,  and  their  posterior  borders  fall  little  behind  the  transverse  line  dividing 
the  skull  equally.  The  interorbital  width  equals  the  long  diameter  of  the  orbit.  The 
posterior  outline  of  the  cranium  is  truncate  in  a  straight  transverse  line  between  the 
prominent  tabulare  angles.  The  composition  of  the  cranium  is  different  from  that  of 
any  other  species  of  Erpetosaurus  in  the  large  size  of  the  tabulare  and  the  fact  that 
the  supratemporal  is  excluded  from  the  parietal  by  the  extension  of  the  postorbitals 
and  the  tabulare.  This  may  be  a  generic  character  and  entitles  the  species  to  be 
placed  in  a  new  genus,  but  it  will  be  retained  here  until  more  of  the  anatomy  of  the 


PLATE   13 


Dendrerpeton  mveni  Dawson.  Above:  Skull,  mandible,  and  bones  of  anterior  limbs.  Below:  Pos- 
terior limb,  pelvis,  and  bony  scales.  Nearly  natural  size.  Erect  tree,  South  Joggins,  Coal 
formation,  Nova  Scotia.  Photograph  by  Dawson,  published  through  the  courtesy  of  Dr. 
Arthur  Willey.     Original  specimen  in  the  Peter  Redpath  Museum  of  McGill  University. 


THE    MICROSAURIAN   FAMILY   TUDITANID^E.  IOI 

species  is  known.  The  elements  of  the  anterior  part  of  the  skull  are  not  preserved, 
but  they  are  indicated  by  the  broken  lines  in  the  drawing  (fig.  22,  G) .  The  nostrils  are, 
however,  clearly  indicated  as  bosses  of  shale.  There  is  a  mere  fragment  of  the  nasal 
preserved  posterior  to  the  crack  indicated  by  the  transverse  line  in  the  drawing. 
The  frontal  is  elongate  as  in  other  species  of  the  genus  and  forms  the  inner  border  of 
the  orbit.  The  parietal,  as  usual,  is  one  of  the  larger  bones  of  the  skull  roof  and  the 
pineal  foramen  is  inclosed  in  the  median  suture  by  the  two  parietal  elements.  The 
pineal  opening  lies  in  the  posterior  half  of  the  parietal.  The  postparietal  is  almost 
square,  being  slightly  elongate  transversely,  uniting  laterally  with  the  tabulare,  with 
which  it  forms  the  truncate  table  of  the  skull.  The  suture  separating  the  tabulare 
from  the  supratemporal  is  clearly  distinct.  Although  such  a  position  for  the  supra- 
temporal  is  unusual  it  is  not  unique,  since  the  same  character  has  been  observed  in 
Diceratosaurus  Icevis  Moodie,  described  elsewhere  (p.  120)  in  this  paper.  The  post- 
frontal  is  rather  small  and  it,  together  with  the  postorbital,  forms  the  posterior 
boundary  of  the  orbit.  The  postorbital  is  truncate  posteriorly  and  joins  the  tabu- 
lare broadly.  The  supratemporal  lies  posterior  to  the  postorbital  and  jugal  and 
borders  the  quadratojugal,  which  is  an  unusual  condition,  but  what  significance 
the  condition  has  remains  to  be  determined.  Posterior  to  the  supratemporal  lies 
the  squamosal,  which  forms  the  quadrate  angle  of  the  skull.  The  quadratojugal  is 
a  small  element  and  forms  part  of  the  lateral  boundary  of  the  skull.  The  jugal  is  a 
large  element  and  forms  the  entire  lateral  border  of  the  orbit.  There  are  no  teeth 
preserved  on  the  fragment  of  the  maxilla,  but  there  are  some  impressions  farther 
forward  which  resemble  the  pleurodont  denticles  of  the  modern  Amphibia. 

The  sculpture  of  the  surface  of  the  cranial  bones  consists  of  parallel  ridges  which 
are  separated  by  grooves  equal  to  them  in  width.  The  ridges  radiate  inward  on  the 
squamosals  and  frontals  and  outward  on  the  supratemporals.  They  are  somewhat 
interrupted  on  the  other  skull  elements.  The  right  clavicle  is  ornamented  with  a 
sculpture  of  similar  radiating  grooves  and  ridges. 

Cope  described  an  atlas  in  connection  with  this  skull,  but  I  do  not  find  it.  The 
slender  impressions  to  the  right  of  the  clavicle  may  possibly  represent  ribs.  They 
are  gently  curved  and  truncate  at  the  inner  end. 

A  nearly  complete  system  of  lateral -line  canals  has  been  detected  on  this  skull. 
The  canals  preserved  are :  the  temporal,  the  jugal,  the  infraorbital,  the  occipital  cross- 
commissure,  and  the  supraorbital.  These  terms  were  used  for  the  first  time  for  the 
Amphibia  by  the  writer  (458)  in  a  discussion  of  the  organs  and  their  significance  in  the 
correlation  of  the  skull  elements.  The  occipital  cross-commissure  in  the  present 
skull  is  represented  by  a  row  of  elongate  pits,  such  as  Andrews  (8)  has  described 
for  Ceraterpeton  galvani  Huxley  from  the  Coal  Measures  of  England.  The  cross- 
commissure  is  contained  within  the  tabulare.  The  jugal  and  temporal  canals  form 
a  complete  ring,  much  as  the  same  canals  do  in  Trematosaurus.  The  supratemporal 
in  Erpetosaurus  tabulatus  Cope  is  excluded  from  the  parietal  by  the  extension  of  the 
tabulare  and  the  postorbital,  and  it  is  to  be  noticed  that  the  temporal  canal  has  a 
changed  position  to  correspond  with  the  changed  condition  of  the  squamosal.  This 
is  of  considerable  interest  in  connection  with  the  correlation  of  the  supratemporal 


102  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

in  fishes  and  amphibians.  This  subject  has  been  fully  treated  in  another  place  (458) 
and  it  will  only  be  necessary  to  state  here  that  on  the  basis  of  the  lateral-line  canals 
and  their  arrangement  in  fishes  and  the  Amphibia  the  true  correlation  of  the  supra- 
temporal  elements  in  amphibians  and  fishes  has  been  made.  The  temporal  canal 
in  the  present  specimen  has,  apparently,  an  indication  of  a  connection  with  the 
supraorbital  canal,  but  of  this  I  am  not  sure.  The  jugal  canal  occurs  on  the  supra- 
temporal  and  quadratojugal,  and  it  joins  the  infraorbital  on  the  jugal.  The  infra- 
orbital is  indicated  by  a  short  portion  a  few  millimeters  long  under  the  orbit  and  the 
remainder,  i.e.,  its  connection  with  the  jugal  canal,  is  restored  (fig.  22,  G).  There  is, 
nothing  unusual  to  be  observed  in  that  portion  of  the  infraorbital  canal  which  is 
preserved.  The  supraorbital  canal  is  indicated  by  a  curved,  broad,  shallow  groove 
on  the  inner  side  of  each  orbit.  As  stated  above,  there  seems  to  be  a  connection 
between  this  canal  and  the  temporal,  but  I  am  not  sure.  The  primitive  conditions 
shown  in  the  lateral-line  canals  in  Erpetosaurus  tabulatus  Cope  are  the  presence  of 
the  occipital  cross-commissure  and  the  ring-like  formation  of  the  temporal  and 
jugal  canals,  which  is  too  clearly  indicated  to  be  overlooked. 

Measurements  of  the  Type  of  Erpetosaurus  tabulatus  Cope. 

mm.  mm. 

Median  length  of  skull 29  Diameter  of  nostril — I 

Width  of  skull,  posterior  border,  estimated 37  Diameter  of  pineal  foramen — 1 

Width  between  tabulare  angles 18  Length  of  right  clavicle 13 

Length  of  orbit 8  Width  of  right  clavicle S-5 

Width  of  orbit 6.5 

MANDIBLE    PROVISIONALLY   ASSOCIATED    WITH    ERPETOSAURUS   TABULATUS    COPE. 
Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  pp.  351-352,  pi.  lix,  fig.  2;  pi.  lxiv,  fig.  3,  1909. 

This  specimen  is  preserved  almost  completely  on  a  slab  of  soft  coal.  It  is  im- 
possible to  determine  positively  to  what  microsaurian  species  the  mandible  be- 
longs, but  it  may,  for  the  present,  be  associated  with  Erpetosaurus  tabulatus  Cope 
on  account  of  its  size  and  the  character  of  its  sculpture.  This  is  the  first  and  almost 
the  only  known  example  of  a  mandible  of  an  American  microsaurian.  The  form  of 
the  jaw  is  perfectly  preserved,  although  the  condition  of  the  articular  surface  can 
not  be  determined. 

The  proportions  of  the  mandible ,  as  may  be  j  udged  from  the  table  of  measurements , 
are  rather  stout  and  the  teeth  are  strong  and  numerous.  There  are  evidences  of  19 
teeth  preserved.  The  sutures  separating  the  articular  (art) ,  angular  (ang) ,  surangular 
(5a) ,  coronoid  (cor) ,  and  the  dentary  (d)  (fig.  22,  F)  are  clear  for  at  least  the  greater  part 
of  their  length  and  they  may  be  easily  restored  for  the  remainder  of  their  course. 
The  surangular  is  thus  seen  to  rival  the  dentary  in  size  and  on  it  occurs  the  peculiar 
sculpturing  which  approximates  so  closely  that  on  the  skull  of  Erpetosaurus  tabu- 
latus Cope.  The  presence  of  the  long  anterior  tooth  is  strikingly  characteristic  of 
many  Microsauria.  It  is  well  developed  in  Sauropleura  longidentata  Moodie  and 
Sauropleura  (Anisodexis)  enchodus  Cope.  It  is  also  present  in  well-developed  form 
in  the  later  labyrinthodonts.  The  teeth  are  all,  with  the  exception  of  the  fourth 
from  the  anterior  end,  rather  short,  curved,  and  sharply  pointed,  with  an  indication 


THE   MICROSAURIAN   FAMILY   TUDITANIOE.  103 

of  longitudinal  fluting.     The  arrangement  of  the  mandibular  elements  recalls  in  a 
striking  way  the  mandible  of  Eryops  megacephalus  Cope  as  figured  by  Branson  (49). 

Measurements  of  the  Mandible  Provisionally  Associated  with  Erpetosaurus  tabulatus  Cope. 
(No.  8542  G,  American  Museum  of  Natural  History.) 

mm.  mm. 

Length  of  mandible 32  Length  of  one  of  the  posterior  teeth 1.25 

Pi  1  terior  width  across  surangular 6  Width  across  base  of  same  tooth 50 

Width  of  dentaiy 3  Length  of  long  anterior  tooth 2 

Width  of  jaw  at  tip 1.5       Width  of  same  tooth  at  base 75 

Another  specimen  of  this  same  species  is  8550  G,  of  the  American  Museum. 
Linton,  Ohio,  Coal  Measures. 

PALATE    OF    ERPETOSAURUS   TABULATUS    COPE. 
Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  pp.  352-354,  pi.  lxi,  fig.  2,  1909. 

The  specimen  is  half  a  cranium  with  its  impression.  It  is  referred  to  Erpeto- 
saurus on  the  basis  of  the  sculpturing  of  the  mandible  and  the  posterior  table  of  the 
skull.  On  the  surangular  there  is  seen  the  rugosity  which  is  common  to  other  mem- 
bers of  the  genus.  The  characters  presented  are  those  of  Erpetosaurus  tabulatus 
Cope,  but  the  reference  is  rather  uncertain. 

The  specimen  is  unique  among  the  known  American  material  in  showing  the 
structure  of  the  palate.  Jaekel  (347)  has  figured  and  described  the  palate  of 
Diceratosa  urus  puuctolineatus  Cope  from  the  Linton  beds.  The  present  palate  differs 
from  that  species  only  in  the  enlarged  ectopterygoid  and  the  smaller  palatine. 

The  parasphenoid,  in  the  present  form,  does  not  differ  from  that  element  in  other 
Paleozoic  Amphibia.  Its  form  is  slender,  arising  from  an  enlarged  base  and  sepa- 
rating the  pterygoids  by  its  own  width.  The  exoccipitals  are  probably  represented  in 
the  specimen  and  they  have  been  indicated  in  the  drawing  (fig.  22  C).  They  are 
rather  large  and  extend  some  distance  under  the  base  of  the  skull  to  unite  anteri- 
orly with  the  pterygoids,  a  very  unusual  arrangement.  The  pterygoids  are  elongate 
elements  and  are  bounded  anteriorly  by  the  vomer  and  laterally  by  the  ectoptery- 
goid. The  vomer  shows  no  evidence  of  being  toothed,  although  it  may  have  been  so 
anteriorly.  The  same  may  also  be  said  for  the  palatines.  The  relations  of  the  ecto- 
pterygoids  are  rather  unusual  for  the  Amphibia,  especially  in  the  posterior  exten- 
sion of  the  element.  The  bone  lies  all  along  the  side  of  the  pterygoid  and  anteriorly 
projects  forward  between  the  pterygoid  and  the  palatine.  In  this  unusual  posterior 
projection  the  ectopterygoid  has  almost  obliterated  the  infratemporal  foramen, 
which  possibly  may  be  still  represented  by  the  triangular  space  between  the  bases 
of  the  pterygoid  and  the  ectopterygoid.  The  anterior  palatine  foramen  (internal 
nares)  lies  between  the  anterior  ends  of  the  palatine  and  the  vomer,  its  usual  rela- 
tions in  the  labyrinthodonts.  The  foramen  may  be  recognized  as  the  rounded 
depression  slightly  anterior  to  the  palatine. 

The  mandible  is  rather  heavy  and  is  coarsely  sculptured  with  radiating  grooves 
and  ridges.  The  character  of  the  teeth  can  not  be  determined,  save  to  say  that  they 
were  present.  The  posterior  end  of  the  mandible  projects  somewhat  beyond  the 
quadrate  angle  of  the  skull. 


104  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

The  interest  in  the  present  specimen  is  heightened  by  the  light  it  throws  on  the 
characters  for  the  separation  of  the  Amphibia  and  Reptilia.  The  wide  separation 
of  the  pterygoids  by  the  parasphenoid  is  an  amphibian  character  of  undoubted  value. 
The  reduction  of  the  parasphenoid  in  this  specimen  is  noteworthy. 

Measurements  of  the  Skull  of  Erpetosaurus  tadulatus  Cope. 
(No.  8607  G,  American  Museum  of  Natural  History.) 

mm.  mm. 

Length  of  skull  in  median  line 45      Width  of  pterygoid 5 

Estimated  posterior  width  of  skull 50      Length  of  ectopterygoid 17 

Estimated  width  of  parasphenoid 6      Posterior  width  of  mandible 12 

Erpetosaurus  minutus  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  pp.  21-23,  pi.  8,  fig.  1,  1909. 

Type:  Specimen  No.  4545,  U.  S.  National  Museum. 

Horizon  and  locality:  Cannelton  slates,  Pennsylvania  (Upper  Freeport). 
(Plate  20 D.) 

The  specimen  on  which  the  species  is  based  is  composed  of  the  greater  portion 
of  a  small  skull  preserved  in  the  hard  shale  from  Cannelton,  Pennsylvania,  and  was 
collected  by  Mr.  R.  D.  Lacoe,  of  Pittston,  Pennsylvania.  The  characters  of  the 
specimen  had  not  previously  been  determined,  since  the  museum  label  and  number 
had  partially  obscured  the  snout  of  the  skull.  The  skull  is  very  small,  but  has  the 
form  assumed  by  other  members  of  the  genus.  The  specimen  may  belong  to  a  young 
individual,  but  even  though  it  does,  it  is,  nevertheless,  quite  distinct  from  the  other 
species  of  Erpetosaurus.  At  first  sight  the  specimen  looks  like  a  broken  scute  of 
some  larger  form.  Close  inspection,  however,  revealed  the  two  impressions  repre- 
senting the  orbits,  and  a  Zeiss  binocular  revealed  the  characters.  The  large  size 
and  anterior  position  of  the  orbits,  the  character  of  the  sculpturing,  the  presence  of 
the  posterior  table  of  the  skull,  as  in  Erpetosaurus  (Tuditanus)  tabulatus  Cope,  are 
the  characters  on  which  a  specific  diagnosis  is  possible.  The  specific  characters 
which  distinguish  this  form  from  E.  tabulatus  Cope,  its  nearest  ally,  are  the  slight 
development  of  the  posterior  table,  the  more  delicate  form  of  the  sculpturing,  the 
more  posterior  position  of  the  orbits,  and  the  varying  shape  assumed  by  the  parie- 
tals  in  the  two  species.  Any  one  of  these  characters  would  be  valid  as  a  specific 
character. 

The  pineal  eye  is  indistinct,  but  is  observed  to  lie  in  the  broken  tract  in  the 
median  line  of  the  skull,  in  the  middle  of  the  portion  posterior  to  the  orbits.  The 
interorbital  space  is  equal  to  the  width  of  the  orbit.  The  orbits  themselves  are 
slightly  oval  and  not  round,  as  in  the  case  of  E.  tabulatus  Cope. 

The  skull  elements  are  sculptured  with  radiating  grooves  and  ridges,  and  on  the 
postparietals  and  tabulare  the  grooves  take  the  form  of  pits  in  a  row,  which  undoubt- 
edly represent  the  occipital  cross-commissure  of  the  lateral-line  system  first  observed 
in  a  microsaurian  by  Andrews  (8)  in  the  skull  of  Ceraterpeton  galvani  Huxley.  The 
supraorbital  canal  is  represented  by  a  slight  elongate  depression  observable  over 
each  orbit  and  extending,  in  one  case,  for  about  5  mm.  The  presence  of  the  circular 
arrangement  of  the  lateral-line  canals  in  the  jugal  region  is  suggested  by  a  depres- 
sion on  the  posterior  edge  of  the  squamosal. 


THE    MICROSAURIAN   FAMILY   TUDITANIDiE.  IO5 

The  portion  of  the  skull  anterior  to  the  orbits  is  wanting,  curiously  enough,  just 
as  in  Erpetosaurus  tabula t us  Cope.  In  the  remainder  of  the  skull,  the  post-parietals, 
the  tabulare,  the  parietals,  the  supratemporal,  and  a  portion  of  the  right  frontal  can 
be  detected,  although  the  boundaries  of  but  three  can  be  accurately  denned.  The 
depression  bounding  the  anterior  outline  of  the  skull  is  taken  to  be  the  impress  of 
the  mandible,  in  which  case  this  structure  would  be  of  some  depth,  as  in  the  case  of 
the  mandible  associated  with  E.  tabulates  Cope,  described  below. 

This  specimen  is  of  interest  in  respect  to  the  presence  of  the  lateral-line  canals, 
its  small  size,  and  its  generic  identity  with  forms  from  Ohio.  There  is  still  another 
form  known  from  the  Cannelton  slates,  described  below  as  Erpetosaurus  (Tuditanus) 
sculpt  His  Moodie. 

Measurements  of  the  Type  of  Erpetosaurus  minutus  Moodie. 

mm.  mm. 

Length  of  skull 18      Length  of  orbit 4.5 

Posterior  width  of  skull 17      Width  of  orbit 3.5 

Width  of  skull  across  orbits 14      Interorbital  width 3.5 

Erpetosaurus  sculptilis  Moodie. 

Moodie,  Jour.  Geol.,  17,  No.  1,  p.  6i,  figs,  n,  12,  1909  {Tuditanus  sculptilis). 
Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  p.  347,  1909  (Erpetosaurus  sculptilis). 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  22,  1909  (describes  pectoral  girdle). 

Type:  Specimen  No.  12,315,  University  of  Chicago,  Walker  Museum. 

Horizon  and  locality:  Cannelton  slates,  Pennsylvania  (Upper  Freeport). 
(Plate  18.) 

There  is  preserved  in  the  collections  of  Walker  Museum  of  the  University  of 
Chicago  a  small  amphibian  skull  pressed  flat  on  a  slab  of  slate  from  Cannelton, 
Pennsylvania.  It  formed  part  of  the  Hall  collection  acquired  by  the  University  of 
Chicago  in  1908. 

The  specimen  presents  only  a  portion  of  the  skull  and  fragmentary  pectoral 
plates.  The  skull  is  wider  than  long  and  the  muzzle  is  broadly  rounded.  The  orbits 
are  narrow  ovals  and  their  posterior  border  falls  on  the  transverse  line  dividing  the 
skull  equally.  The  interorbital  width  is  slightly  greater  than  the  width  of  the  orbits 
and  about  equal  to  their  length.  The  posterior  outline  of  the  skull  is  somewhat 
truncate,  as  in  E.  tabulates  Cope  and  other  species  of  the  genus.  The  distal 
extremities  of  the  quadrates  do  not  project  as  far  backward  as  do  the  supraoccipitals. 
The  skull  roof  is  formed  of  the  regular  elements,  except  that  a  quadrate  seems  to  be 
indicated  by  a  scale  of  bone  on  the  posterior  angle.  The  nostrils  are  oval  and  the 
pineal  opening  is  small. 

The  premaxillae  are  apparently  relatively  large  elements,  though  their  bounda- 
ries are  not  definite.  The  nasal  is  of  an  oblong  shape  and  borders  the  frontal  ante- 
riorly. The  frontal  forms  the  whole  of  the  interior  border  of  the  orbit  and  borders 
the  parietal  broadly  behind.  The  parietal  is  a  large  element  and  the  pineal  foramen 
is  inclosed  in  the  median  suture  about  midway  of  the  parietals.  The  postparietal 
is  wider  than  long  and  with  the  tabulare  forms  the  greater  part  of  the  posterior 
border  of  the  skull.  The  prefrontal  (plate  18,  fig.  1)  apparently  forms  the  entire 
anterior  border  of  the  orbit  and  sends  an  acuminate  projection  to  the  side  of  it.    The 


IOG  THE    COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

maxilla  is  excluded  from  the  orbit  and  is  an  elongate  element  with  sharp  conical 
teeth,  of  which  there  are  4  preserved.  These  measure  about  1  mm.  in  length.  The 
jugal  lies  along  the  lateral  border  of  the  orbit  and  it  is  acuminate  both  anteriorly 
and  posteriorly.  It  borders  the  supratemporal  broadly.  The  postfrontal  forms  the 
greater  part  of  the  posterior  boundary  of  the  orbit.  It  is  triangular  and  acuminate 
behind,  and  is  bordered  broadly  by  the  parietal  and  supratemporal.  The  supratem- 
poral is  also  triangular  and  it  borders  the  parietal  broadly.  The  squamosal  is  evi- 
dently the  largest  element  in  the  skull  and  on  its  posterior  corner  there  is  a  flake  of 
bone  which  may  represent  the  quadrate,  though  this  is  by  no  means  certain.  The 
quadrate  has  not  been  detected  in  any  of  the  Carboniferous  Microsauria  so  far 
studied.  The  tabulare  is  an  elongate  element  in  the  transverse  line  of  the  skull. 
Its  entire  boundary  is  uncertain,  though  part  of  the  sutures  are  present.  The 
quadratojugal  is  elongate  and  lies  posterior  to  the  maxilla  and  with  that  element 
forms  the  lateral  boundary  of  the  skull. 

The  canals  of  the  lateral-line  system  have  not  been  detected  on  the  skull.  The 
sculpturing  of  the  cranial  elements  consists  of  grooves  and  ridges  which  radiate  from 
a  center.  They  are  more  prominent  on  the  parietals  than  elsewhere,  although  the 
other  skull  elements  present  a  strong  sculpturing. 

There  are  also  preserved  on  the  slab  of  slate,  about  10  mm.  posterior  to  the  skull, 
fragments  of  the  pectoral  plates,  probably  representing  the  clavicles  and  the  inter- 
clavicle.  They  are  so  badly  fractured  that  their  form  can  not  be  determined.  No 
limbs  or  vertebrae  have  been  observed. 

Measurements  of  the  Type  of  Erpetosaurus  sculptilis  Moodie. 

mm.  mm. 

Length  of  skull  in  median  line 20  Interorbital  space 4 

Width  of  skull  at  posterior  border  (estimated) 24  Diameter  of  nostril — 1 

Diameter  of  orbit 3  Pineal  foramen,  diameter 50 

Length  of  orbit 4 

Pectoral  Girdle  Provisionally  Associated  with  Erpetosaurus  sculptilis  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  22,  1909. 

The  present  specimen  is  preserved  on  a  block  of  slate  from  Cannelton,  Pennsyl- 
vania. It  is  associated  with  the  previously  described  Erpetosaurus  sculptilis  Moodie 
on  account  of  its  size,  the  geological  and  geographical  distribution,  and  the  charac- 
ter of  the  sculpture.  It  may  pertain  to  an  unknown  species.  Other  remains  besides 
the  3  elements  of  the  pectoral  girdle  are  preserved  on  the  block  of  slate,  but  they 
are,  for  the  most  part,  too  imperfectly  preserved  for  recognition.  Some  of  them  are 
phalanges,  and  I  believe  I  detect  a  scapula  in  the  rounded  curved  plate  lying  near 
the  right  clavicle.  The  3  pectoral  elements,  the  interclavicle  and  the  2  clavicles,  are 
preserved  intact,  with  the  ventral  surface  uppermost. 

The  specimen  is  particularly  important  in  that  it  furnishes  further  evidence  of  the 
simplicity  of  the  microsaurian  pectoral  girdle,  which  Jaekel  regarded  (347)  as  being 
extremely  complex,  in  one  species  at  least,  Diceratosaurus  punctolineatus  Cope. 
The  3  elements  are  broken,  but  either  the  elements  or  their  impressions  are  pres- 
ent, so  that  identification  is  possible.     The  elements  are  sculptured  with  radiat- 


1  and  2.     Photograph  of  the  specimen  of  Amphibamus  grandiceps  Cope,  from  the  Mazon  Creek   shales. 
X  1.5.     Original  in  possession  of  Mr.  L.  E.  Daniels,  Rolling  Prairie,  Indiana. 

3.  Specimen  of  Sauropleiira  (Coloslnis)  saitrllata  Newberry,  from  the  Linton  Coal  Measures.  The  species 

was  the  first  known  of  the  Ohio  Coal  Measures  Amphibia;  at  first  ascribed  by  Newberry 
to  the  fishes,  but  later  correctly  identified  by  Cope.  X  1.  Original  in  American  Museum 
of  Natural  History. 

4.  The  type  of  Diceratosaurus  (CeraterpetoiA  punctolineafus  Cope,  from  the  Linton  Coal  Measu res.     X  1. 

Original  in  American  Museum  of  Natural  History. 


THE   MICROSAURIAN   FAMILY  TUDITANID^E.  107 

ing  grooves  and  ridges,  as  in  so  many  of  the  Microsauria.  The  interclavicle  is 
spatulate  and  bears  a  general  resemblance  to  the  same  element  of  Metoposaurus 
fraasi  Lucas  from  the  Triassic  (383)  of  Arizona.  The  clavicles  are  triangular, 
with  rounded  angles  and  the  hypothenuse  on  the  interior  border. 

Measurements  of  Pectoral  Girdle  of  Specimen  No.  4539,  U.  S.  National  Museum. 

mm.  mm. 

Width  across  the  entire  girdle 17      Length  of  clavicle 11 

Length  of  interclavicle 15      Width  of  clavicle,  maximum 6 

Width  of  interclavicle 10 

Erpetosaurus  acutirostris  Moodie. 
Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  pp.  349-351,  pi.  lxi,  fig.  1,  1909. 

Type:  Specimen  No.  8598  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  present  species  adds  another  form  to  the  diversity  of  structure  presented  by 
the  Carboniferous  Microsauria.  It  is  closely  allied  to  Erpetosaurus  (Tuditanus)  obtu- 
sus  Cope,  from  the  same  beds  (Linton,  Ohio),  but  differs  from  it  especially  in  the 
position  and  shape  of  the  orbits  and  the  acute  form  of  the  skull.  Other  characters 
which  amount  almost  to  generic  significance  are  found  in  the  posterior  prolongation 
of  the  frontal  and  in  the  triangular  form  of  the  skull.  Only  the  skull  of  the  animal 
is  preserved.  The  character  which  is  common  to  all  members  of  the  genus  Erpeto- 
saurus, the  cranial  rugosity,  is  present  in  this  species  on  the  squamosal  and  supra- 
temporal.  This  character  alone  would  not,  however,  suffice  to  separate  the  form 
generically,  but  the  general  morphology  and  arrangement  of  the  cranial  elements  is 
such  that  reference  to  any  other  genus  save  Erpetosaurus  would  not  be  possible. 

The  skull  of  E.  acutirostris  takes  the  form  of  a  rounded  triangle.  Its  base  is 
some  50  mm.  in  extent,  and  this  width  gradually  narrows  to  31  mm.  across  the 
orbits  and  still  more  towards  the  snout.  The  form  of  the  skull  is  not  widely 
different  from  that  of  the  type  species,  E.  radiatus  Cope,  but  the  differences  are 
sufficiently  apparent. 

Nearly  all  the  elements  of  the  cranium  can  be  detected  (fig.  22,  D).  The  bony 
portion  of  the  cranium  has  nearly  all  been  lost,  leaving  only  the  impression;  and 
the  matrix  in  which  the  skull  was  embedded  has  been  forced  up  into  the  sutures 
between  the  cranial  elements,  thus  forming  ragged  ridges  where  the  bones  of  the 
skull  joined. 

The  position  of  the  nostrils  can  not  be  determined  accurately.  The  orbits  are 
placed  well  forward,  a  character  common  to  several  species  of  the  genus.  The  inter- 
orbital  space  is  equal  to  the  long  diameter  of  the  eye.  The  orbits  are  sep- 
arated by  narrow  prolongations  of  the  postfrontals  and  by  the  anterior  por- 
tion of  the  frontals.  The  frontals  are  remarkable  in  their  great  back- 
ward extension.  In  E.  obtusus  the  frontals  are  nearly  confined  to  the  inter- 
orbital  space.  The  parietals,  which,  on  the  median  suture,  inclose  the 
parietal  foramen,  lie  well  posterior,  and  the  parietals  and  the  tabulare  are 
small.     A  portion  of  the  sculpturing  of  these  elements  has  been  preserved  and 


108  THE    COAL   MEASURES    AMPHIBIA   OF   NORTH    AMERICA. 

it  is  seen  to  be  made  up  of  pits  and  elevations  much  as  we  find  in  the  skull  of 
Saurerpeton  latithorax  Cope.  The  left  squamosal  also  shows  sculpturing,  which  here 
tends  to  take  the  form  of  grooves  and  ridges,  and  also  of  pits  and  elevations.  It  is 
quite  probable  that  the  anterior  portion  of  the  skull  was  ornamented  with  grooves 
and  ridges  and  undoubtedly  the  lateral-line  canals  were  well  developed.  The  post- 
frontals  and  the  postorbitals  are  both  large  and  elongated.  The  postorbital  is 
especially  large.  The  supratemporal  apparently  separates  the  tabulare  and  the 
squamosal  in  their  posterior  extremities.  The  squamosal  projects  posteriorly  to  the 
tabulare  and  apparently  even  goes  beyond  the  limits  of  the  exoccipital.  The  out- 
lines of  the  jugal  are  fairly  definite,  as  are  also  the  limits  of  the  maxilla. 

Measurements  of  the  Type  of  the  Skull  of  Erpetosaurus  acutirostris  Moodie. 

mm.  mm. 

Length  of  skull 50  Width  across  orbits 31 

Interorbital  space 9  Posterior  width  of  skull 50 

Width  of  orbit 7  Diameter  of  pineal  foramen I 

Length  of  orbit 10  Length  from  tip  of  snout  to  posterior  angle  of  skull. .   65 

An  additional  specimen  shows  that  the  skull,  of  which  the  anterior  half  is  pre- 
served, is  practically  of  the  same  size  as  the  type  and  shows  much  the  same  characters, 
though  more  extensively.  The  sculpture  of  the  squamosal  region  is  not  confined  to 
that  portion  of  the  skull,  but  extends  throughout  the  cranial  elements,  apparently 
including  the  premaxillae. 

The  sutures,  which  are  clearly  distinct,  are  of  the  same  type  as  has  been  described 
for  the  type  specimen.  Perhaps  one  of  the  most  interesting  characters  discovered 
on  the  present  specimen  is  that  of  the  greater  part  of  the  left  supraorbital  lateral- 
line  canal,  which  is  exhibited  as  a  rather  deep  and  broad  canal  running  in  a  slight 
curve  across  the  lower  edges  of  the  postorbital  and  the  parietals  and  partly 
cutting  the  jugal. 

On  the  left  of  the  fossil,  as  it  is  preserved,  there  is  an  indistinct  impression  of  the 
clavicle,  with  the  sculpture  in  radiating  lines  from  the  apex  as  a  center,  such  as  have 
been  found  in  other  species  of  the  Tuditanidae. 

Measurements  of  Additional  Specimen. 
(No.  8607,  American  Museum  of  Natural  History.) 

mm.  mm. 

Length  of  skull  as  preserved 40      Length  of  clavicle 18 

Anterior  width  across  orbital  region 22      Greatest  width  of  clavicle 9 

Greatest  width  of  the  skull 33 

Still  a  third  specimen  of  this  species  is  possibly  represented  by  a  nearly  complete 
skull  of  a  small  individual  which  exposes  the  mandible  and  the  ventral  portion  of  the 
skull.  The  remains  are  crushed  flat,  though  not  at  all  distorted.  It  is  quite  pos- 
sible that  the  present  specimen  represents  a  distinct  species,  but  only  a  small  portion 
of  the  dorsum  is  present  and  the  shape  of  the  cranium  is  indistinct,  so  it  is  retained 
in  this  species.  The  portion  of  the  skull  shows  the  sculpture  to  be  quite  similar  to 
that  of  Erpetosaurus  acutirostris,  so  far  as  the  species  is  known;  and  the  skull  appar- 
ently tapers  to  a  point  anteriorly.  It  may  be  a  dwarfed  or  immature  form.  The 
sculpturing  of  the  jaw  is  such  as  we  would  expect  of  this  species. 


THE   MICROSAURIAN   FAMILY   TUDITANIDiE.  109 

The  specimen  is  about  half  the  size  of  the  type.  The  palate  of  the  skull  is  well 
preserved  and  is  extremely  interesting.  The  sutures  separating  the  various  palatal 
elements  are  not  distinct.  The  parasphenoid  is  especially  large  and  the  exoccipitals 
arc  partly  ossified,  if  we  may  judge  by  the  projecting  condyles.  Anteriorly  the 
parasphenoid  contracts  and  then  expands  and  on  each  side  of  the  expanded  part  lie 
fragments  of  the  palatines.  To  the  right  of  the  posterior  end  of  the  parasphenoid 
lies  a  portion  of  the  dorsal  element  showing  the  cranial  sculpture. 

The  left  mandible  is  somewhat  displaced  to  the  right  of  the  skull,  and  crushed 
and  weathered  to  such  an  extent  that  the  sutures  are  entirely  obliterated.  There 
are  3  teeth,  with  indications  of  others.  They  are  typically  pleurodont  and  sharp 
and  slender.  The  mandible  tapers  somewhat  anteriorly  and  at  the  tip  bears  an 
elongate  enlarged  tooth. 

Measurements  of  the  Third  Specimen  of  Erpetosaurus  acutirostris  Moodie  in  the  American 

Museum  of  Natural  History. 

mm.  mm. 

Length  of  skull,  as  preserved 25      Length  of  mandible 25 

Posterior  width  of  skull 15      Anterior  width  of  mandible 2 

Anterior  width  of  skull 10      Length  of  large  tooth I 

Width  across  occipital  condyles 4 

Erpetosaurus  tuberculatus  Moodie. 
Moodie,  Bull.  Amer.  Mus.  Nat.  Hist.,  xxvi,  pp.  348-349,  pi.  lviii,  1909. 

Type:  Specimen  Nos.  8693  G  and  8610  G,  American  Museum  of  Natural  His- 
tory. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  species  is  based  on  a  fragmentary  cranium  (plate  26,  fig.  1)  consisting  of  the 
posterior  part  of  the  right  side  of  the  skull.  Its  association  in  the  genus  is  solely  on 
the  character  of  the  sculpturing  of  the  cranial  elements.  It  is  most  closely  related, 
in  the  characters  preserved,  to  the  form  described  by  Cope  as  Tuditanus  radiatus, 
from  which  it  differs  especially  in  the  character  of  the  sculpture  and  in  the  position 
of  the  orbits,  as  well  as  the  arrangement  and  size  of  the  various  cranial  elements,  so 
far  as  these  elements  can  be  detected  in  the  present  specimen.  In  Erpetosaurus  radi- 
atus the  skull  is  sculptured  by  radiating  grooves  and  ridges  which  did  not  arise  from 
a  definite  center.  In  E.  tuberculatus  this  center  of  radiation  is  marked  by  an  eleva- 
tion or  tubercle  on  each  cranial  element  exposed,  from  which  the  grooves  and  ridges 
radiate  outward.  These  tubercles  have  an  elevation  of  4  mm.  above  the  cranial 
element  proper.  The  orbit  is  located  near  the  median  line  of  the  skull,  so  far  as  can 
be  determined.  In  E.  radiatus  Cope  the  orbits  are  located  well  forward.  In  that 
species  also  the  postparietal  is  smaller  than  in  the  present  species  and  the  squamosal 
is  longer  and  more  slender.     (Plate  25,  fig.  1.) 

The  fragment  of  a  skull  on  which  the  above  comparison  has  been  made  consists 
of  the  right  postparietal,  a  portion  of  the  tabulare,  the  parietal,  the  frontal,  and  a 
portion  of  the  squamosal.  The  other  elements  are  not  clear.  The  elements  in  the 
median  line  are  elongate,  as  in  Erpetosaurus  radiatus.  The  pineal  foramen  is  located 
well  back  on  the  median  line  and  lies  posterior  to  two-thirds  of  the  length  of  the 


HO  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

parietals.    The  sutures  separating  the  frontal  and  parietal  elements  from  each  other 
in  the  median  line  are  of  the  zigzag  form  so  characteristic  of  the  labyrinthodonts. 

Measurements  of  the  Type  of  Erpetosaurus  tuberculatus  Moodie. 

mm.  mm. 

Length  of  portion  of  skull  preserved 52  Maximum  width  of  parietal n 

Width  across  tabulare  (estimated) 60  Length  of  frontal 22 

Length  of  parietal 15  Width  of  frontal 12 

Genus  ODONTERPETON  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  19,  1909. 

Type:  Odonterpeton  triangularis  Moodie. 

The  generic  characters  may  be  found  in  the  triangular  shape  of  the  skull,  the 
large  size  of  the  teeth,  the  shape  of  the  vertebrae,  the  small  size  of  the  orbits,  and 
their  anterior  position  as  shown  in  the  type  specimen  (fig.  22,  E).  The  name  of  the 
genus  is  derived  from  the  remarkable  size  of  the  teeth  as  compared  with  the  size 
of  the  skull. 

Odonterpeton  triangularis  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  19,  pi.  6,  fig.  3,  1909. 

Type:  Specimen  No.  4465,  U.  S.  National  Museum. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  specimen  described  under  the  above  name  is  a  representative  of  the  small- 
est species  of  the  Microsauria  so  far  described  from  North  America.  Orthocosta 
microscopica  Fritsch,  from  the  Carboniferous  of  Bohemia,  is  a  rival  of  the  present 
form  in  size,  but  the  form  described  by  Fritsch  is  an  entirely  different  animal  and 
was  formerly  included  among  the  so-called  Aistopoda,  which  are  regarded  by  the 
writer  as  merely  specialized  microsaurians.  The  present  form  shows  clear  affinities 
with  the  Microsauria. 

As  may  be  seen  by  referring  to  the  list  of  measurements,  the  skull  of  this  form 
measures  only  6.5  mm.  in  length.  The  form  may  possibly  be  larval,  though  I  do 
not  think  so,  if  I  may  judge  from  the  well-developed  condition  of  the  skull  bones 
and  the  complete  ossification  of  the  vertebras.  The  sides  of  the  skull  are  equal  and 
the  occiput  is  a  straight  table,  so  that  the  skull  forms  almost  an  exact  equilateral 
triangle.  The  orbits  are  very  small  and  are  placed  well  forward.  The  interorbital 
space  is  four  times  that  of  the  diameter  of  the  orbit,  a  very  unusual  character  and 
in  itself  worthy  of  ranking  as  a  generic  character.  The  median  suture  of  the  skull 
is  zigzag  and  incloses  the  minute  parietal  foramen  near  the  posterior  end  of  the 
skull.  The  relations  of  the  elements  of  the  skull,  with  the  exceptions  of  those  of 
the  frontals  and  parietals,  can  not  be  determined  with  accuracy,  although  there  are 
here  and  there  indications  of  sutures.  The  characters  of  the  cranial  elements,  so 
far  as  they  can  be  determined,  are  those  of  the  family  Tuditanidae,  and  the  form 
may,  for  the  present,  be  regarded  as  a  member  of  that  group.  The  teeth  are  very 
long,  slender,  and  sharp,  and  are  placed  close  together.  There  is  no  indication  of 
fluting  on  the  teeth.    They  are  slightly  curved  inward. 

There  are  13  vertebrae  present.  The  centra  are  hour-glass  shaped,  and  are 
apparently  phyllospondylous,  with  the  notochord  largely  persistent.     The  vertebral 


THE   MICROSAURIAN    FAMILY   TUIHTANID^.  Ill 

centra  are  unusually  long  and  slender,  with  the  ends  rounded.  The  humerus  of  the 
right  side  is  preserved.  It  is  a  long,  slender  bone  with  expanded  extremities.  There 
is  no  evidence  of  abdominal  armature  nor  of  ribs  (fig.  22,  E). 

The  discovery  of  this  form  in  the  Linton  deposits  is  of  considerable  interest  in 
that  it  indicates  a  wide  range  in  size  and  character  of  the  fauna  of  the  time.  The 
forms  now  known  from  the  Linton  beds  range  from  Odonterpeton,  which  possibly  had 
a  total  length  of  2  inches  in  life,  to  the  form  designated  Macrerpeton  huxleyi  Cope, 
with  a  skull  of  at  least  8  inches  in  length  and  whose  body  may  have  attained  a  length 
of  some  feet.  The  large  rib  described  below  undoubtedly  indicates  a  large  form  of 
the  ancient  Amphibia  from  Linton,  as  do  also  the  vertebrae  described  by  Marsh  in 
1863  from  Nova  Scotia. 

Measurements  of  the  Type. 

mm.  mm. 

Length  of  animal,  as  preserved 18  Length  of  tooth 0.25 

Length  of  skull 6.5  Length  of  vertebra 1.45 

Posterior  width  of  skull 5.5  Width  of  vertebra 35 

Length  of  side  of  skull 6.5  Length  of  humerus 2.25 

Diameter  of  orbit 65  Distal  width  of  humerus 35 

Intcrorbital  width 2 


CHAPTER  XV. 

THE  MICROSAURIAN  FAMILY  STEGOPID^E,  FROM  THE  COAL  MEASURES  OF  OHIO. 

Family  STEGOPIDjE  Moodie,  1909. 

Moodie,  Jour.  Geol.,  xvii,  No.  I,  79,  1909. 

The  chief  family  characters  are  the  large  lacrimal,  unknown  in  other  species 
of  Coal  Measures  Amphibia,  the  central  position  of  the  orbits,  the  general  form  of 
the  skull,  and  the  peculiar,  short,  divaricate  horns  from  the  squamosal.  If  an  inter- 
temporal element  is  present  in  the  skull,  which  is  suggested  as  a  possibility,  the 
family  is  further  distinct.  The  type  species  is  Stegops  divaricata  Cope  from  the 
Linton,  Ohio,  Coal  Measures. 

The  group  seems  to  be  distinct  and  has  no  immediate  allies,  being  confined  to 
the  American  Coal  Measures. 

Genus  STEGOPS  Moodie,  1909. 
Moodie,  Jour.  Geol.,  xvii,  79,  1909. 

Type:  Stegops  divaricata  Cope. 

This  genus  has  been  erected  for  the  reception  of  the  peculiar  form  described  by 
Cope  as  Ceraterpeton  divaricatum,  but  there  are  good  reasons  why  the  form  can  not 
be  retained  in  the  genus.  The  position  of  the  orbits  in  Stegops  (plate  25,  fig.  3)  is  dif- 
ferent from  Ceraterpeton  and  the  muzzle  is  rounded,  not  truncate  as  in  the  latter 
form.  The  horns  are  of  a  different  type  and  there  is  no  indication  of  the  tabulare 
protuberance  which  is  present  in  Ceraterpeton.  The  sculpturing  of  the  cranial 
elements  is  also  distinctive  in  the  present  form,  consisting  of  radiating  grooves  and 
ridges;  the  cranium  of  Ceraterpeton  appears  to  be  but  slightly  sculptured.  There 
is  no  lateral  projection  from  the  border  of  the  skull  in  Stegops,  as  there  is  in  the 
other  genus.  The  structure  of  the  skull  of  Ceraterpeton  is  practically  unknown, 
except  in  a  very  general  way,  although  Andrews  (8)  was  able  to  make  out  some  of 
the  elements  and  to  trace  the  lateral-line  canals.  A  structural  comparison  is  thus 
impossible,  but  on  the  basis  of  form  alone  there  are  good  generic  distinctions.  The 
present  genus  is  apparently  distinct  from  other  genera  in  the  presence  of  an  inter- 
temporal, but  additional  material  will  be  required  before  a  satisfactory  determina- 
tion is  possible.  The  genus  Diceratosaurus  of  Jaekel  (347)  is  distinct  in  the  arrange- 
ment of  the  elements  of  the  cranium,  the  general  form  of  the  skull,  and  in  the  two 
known  species  of  Diceratosaurus  the  orbits  are  located  well  anteriorly,  but  in 
Stegops  they  are  in  the  median  transverse  line  of  the  cranium.  The  genus  Stegops  is 
distinct  from  Eoserpeton  in  the  smaller  size  of  the  prosquamosal,  in  the  broadly 
rounded  muzzle,  in  the  larger  and  more  posteriorly  placed  orbits,  and  in  the  pres- 
ence of  an  intertemporal  bone,  or  at  least  in  the  elongate  character  of  the  post- 
orbital  if  the  intertemporal  is  not  present.  The  species  on  which  the  genus  Eoserpe- 
ton is  based  was  first  described  by  Cope  as  Ceraterpeton  tenuicorne.    The  form  is 


THE   MICROSAURIAN   FAMILY   STEGOPIM;. 


113 


quite  distinct,  in  spite  of  Jaekel's  protestations  to  the  contrary.  The  genus  Stegops 
stands  alone  among  the  Carboniferous  Amphibia  of  North  America,  so  far  as  I  am 
aware,  in  the  possession  of  a  well-defined  lacrimal  of  the  labyrinthodont  type. 

Stegops  divaricata  Cope. 


Cope,  Proc.  Am.  Phil.  Soc.,  xxn,  p.  406,  1885  (Keraterpeton  divaricatum) . 
Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  79,  fig.  22,  1909  {Stegops). 


The 


Type:  Specimen  No.  2559  G,  American  Museum  of  Natural  History 
obverse  of  this  is  No.  12,311,  Walker  Museum,  University  of  Chicago. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  skull  on  which  this  species  is  based  consists  of  the  impressions  on  two  slabs 
of  coal,  one  belonging  to  the  Newberry  Collection  of  the  American  Museum  of 
Natural  History,  No.  2559  G,  and  the  other  to  the  Gurley  Collection  of  the  Uni- 
versity of  Chicago,  No.  12,311.  The  slab  be- 
longing to  the  University  of  Chicago  contains  the 
better-preserved  remains,  so  that  the  descrip- 
tion is  based  largely  on  that  portion  (plate  25, 
fig.  3).  Nearly  all  of  the  elements  of  the  skull 
are  determined  with  considerable  certainty  and 
many  important  characters  in  the  morphology 
of  the  Microsauria  are  thus  brought  out. 

The  skull  is  oval,  elongate,  truncate  behind, 
and  the  quadrate  angles  project  into  sharp 
horns.  The  orbits  are  elongate  ovals  and  their 
center  lies  in  the  median  line  which  divides  the 
skull  transversely.  The  nostrils  are  elongate 
and  have  an  oblique  position.  The  pineal  fora- 
men lies  in  the  posterior  third  of  the  skull. 
Teeth  are  preserved  on  both  maxilla?  and  pre- 
maxillae.  They  are  simply  sharp  pleurodont 
denticles,  and  seem  to  have  been  fairly  abun- 
dant. The  bones  have  been  completely  carbon- 
ized and  nothing  of  the  original  texture  is  preserved,  although  the  details  of  the 
structure  are  beautifully  preserved.    (Plate  25,  fig.  3.) 

The  skull  is  somewhat  triangular  in  its  general  form.  The  premaxilla  lies  on 
the  anterior  border  of  the  cranium,  and  forms  the  median  border  of  the  nostril. 
The  suture  which  separates  the  maxilla  and  the  premaxilla  is  not  evident,  and  it 
may  not  be  correctly  defined  in  the  figure  (fig.  23).  The  nasal  is  a  very  large 
element  and  is  elongate.  It  unites  with  the  premaxilla,  the  lacrimal,  the  pre- 
frontal, and  the  parietal.  It  is  acuminate  behind  and  the  point  is  inclosed  by  the 
prefrontal  and  the  parietal.  The  frontal  is  quite  narrow  and  elongate,  and  does 
not  border  the  orbit;  its  posterior  boundary  is  not  accurately  represented.  The 
radiations  on  the  surface  indicate  the  extent  of  the  element.  The  parietals  are 
remarkable  in  being  smaller  than  the  frontal  and  nasal.     The  pineal  foramen  is 


Fig.  23. — Skull  elements  of  Stegops  divaricata 
Cope,  ft,  frontal ;  it,  intertemporal ;  j,  jugal ; 
la,  lacrimal;  mx,  maxilla;  n,  nasal;  par, 
parietal;  pf,  prefrontal;  po,  postorbital;  pof, 
postfrontal;  pmx,  premaxilla;  sq,  squamosal; 
spt,  supratemporal ;  qj,  quadratojugal;  tab, 
tabulare;  pp,  postparietal.    X  1.3. 


114  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

inclosed  in  the  median  suture  in  the  anterior  third  of  the  parietals.  The  parietal 
has  the  usual  relations  of  that  element.  The  postparietal  lies  posterior  to  the  parie- 
tal, but  a  portion  of  its  bounding  sutures  are  destroyed.  The  prefrontal  forms  the 
antero-interior  border  of  the  orbit  and  borders  the  postfrontal  posteriorly,  in  an 
unusual  manner.  The  lacrimal  is  a  large  element  and  is  clearly  separable  from 
the  other  cranial  elements.  It,  with  the  prefrontal,  forms  the  anterior  border  of 
the  orbit.  The  maxilla  is  very  elongate  and  forms  the  larger  part  of  the  lateral 
border  of  the  skull.  Sharp-pointed  teeth  are  present  on  it  and  they  may  have  been 
pleurodont.  The  lateral  border  of  the  orbit  probably  received  a  portion  of  the 
maxilla.  The  postfrontal  and  the  postorbital  form  the  greater  part  of  the  poste- 
rior boundary  of  the  orbit.  The  postorbital  seems  to  be  divided  by  a  median  suture 
which  would  indicate  an  intertemporal  bone,  but  this  is  not  certain.  The  appear- 
ance may  be  due  to  a  fracture.  The  supratemporal  is  a  large  element  bordering 
the  parietal  and  lies  in  front  of  the  tabulare.  The  squamosal  is  elongate  to  form 
the  posterior  horn -like  extension  of  the  skull.  The  tabulare  is  transversely  elon- 
gate and  has  the  usual  relations.  The  jugal  widens  to  a  fan-shape  backwards,  and 
helps  to  form  the  lateral  border  of  the  orbit.  Its  lateral  and  anterior  boundaries 
are  not  assured.  The  quadratojugal  seems  to  lie  as  indicated,  although  the  ante- 
rior part  of  the  suture  is  not  distinct.  It  is  apparently  an  elongate  element  and  with 
the  maxilla  forms  the  lateral  border  of  the  skull.  The  base  of  the  skull  as  restored 
(fig.  23)  is  irregular  and  may  have  had  a  slightly  different  form. 

The  genus  Stegops  is  exceptional  in  the  elongate  character  of  the  cranial  ele- 
ments of  the  single  species  known.  In  this  respect  it  recalls  the  species  Dicer ato- 
saurus  Icevis  described  below.  The  large  size  of  the  nasals,  frontals,  and  lacrimals 
and  the  small  size  of  the  parietals  are,  so  far  as  I  am  aware,  unparalleled  among  the 
other  Coal  Measures  Amphibia  of  North  America. 

Measurements  of  the  Type. 


mm. 


Median  length  of  the  skull 56  Interorbital  space 16 

Width  across  tips  of  horns 46  Length  of  the  nostril 2 

Width  at  base  of  horns 40  Diameter  of  the  pineal  foramen 1 

Width  across  orbits 44  Length  of  the  teeth 1.5 

Diameter  of  the  orbit 8  Length  of  the  horn  from  base 7.5 

Length  of  the  orbit 15  Width  of  horn  at  base 4 


MOODIE 


1.  Dorsum  of  skull  of  Diceratosaurns   punctolineatus   (Cope),    from   the    Coal    Measures    of   Linton, 

Ohio.  Original  in  the  Museum  at  Berlin  University.  X  2.  After  Jaekel.  /o=postorbital; 
/r=frontal;  ^o/=postfrontal;  ;'=jugal;  /a=laerimal;  »/j-=maxilla;  /».s="perisquamosal;" 
pp= post-parietal;  ^«/:r=premaxilla. 

2.  Ventral  surface  of  the  skull  of  Diceratosaurns punctolineatus  (Cope),  from    the   Coal    Measures   of 

Linton,  Ohio.  X  2.  After  Jaekel.  «=anterior  palatine  vacuity;  *r=:exoccipital;  /=jugal; 
»tr=maxilla;/r=transverse;/><7/=palatine;^=pterygoid;^r=prevomer;/.s=''perisquamosal;'' 
^z'=posterior  or  suborbital  palatine  vacuity;  /A=parasphenoid. 

3.  Pectoral  girdle  of  Diceratosaurns  punctolineatus  (Cope),  from  the  Coal  Measures  of   Linton,   Ohio. 

Original  in  the  paleontological  Museum  at  Berlin.  X  2.  After  Jaekel.  f//'=inner  side  of 
clavicle;  ;'r=interclavicle;  <7=clavicle  (.lower  side);  fte=cleithrum. 

4.  (e )  Cervical  or  anterior  dorsal  vertebra  of  Diceratosaurns  punctolineatus  (Cope),  from  the   Linton, 

Ohio,  Coal  Measures.  rj<=c:uidal  vertebra  shown  with  its  cap  of  dermal  bone;  dv—<\oxs&\ 
vertebra  with  ribs;  .r-dcrmal  plate  <>n  neural  spine;  A=humeius;  «=ulna;  r—  radius. 


CHAPTER  XVI. 

THE  MICROSAURIAN  FAMILY  UROCORDYLID^E,  FROM  THE  COAL  MEASURES 

OF  OHIO. 

Family  UROCORDYLID^E  Lydekker,  1890. 

L<i  DECKER,  Cat.  Fossil  Rcptilia  and  Amphibia,  pt.  iv,  p.  196,  1890. 
Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  art.  xxv,  p.  357,  1909. 

Type  of  the  family:   Urocordylus  wandesjordii  Huxley. 
Locality  and  horizon:  Coal  Measures  of  Kilkenny,  Ireland. 

Stout  and  long-tailed  forms,  with  the  tabulare  cornua  frequently  much  pro- 
duced and  pitted  cranial  bones;  lateral  lines  well-developed;  palate  with  teeth  on 
palatines,  vomers,  premaxillas,  and  maxillae,  the  two  latter  elements  bearing  conical 
teeth  and  the  others  bearing  short,  stumpy  cones,  at  least  in  one  species;  pineal 
foramen  well  forward;  nostrils  and  orbits  in  the  anterior  part  of  skull;  scapula 
peculiarly  curved  and  pointed;  other  pectoral  elements  sculptured;  neural  spines 
and  chevrons  of  caudal  vertebrae  much  dilated  at  their  extremities,  and  pectinated ; 
no  caudal  ribs ;  vertebra  in  one  genus  apparently  capped  with  a  sculptured  plate 
as  in  Zatrachys;  tail  very  long  and  tapering  to  a  point,  50  to  80  caudal  vertebrae; 
dorsal  region  short;  limbs  well  developed,  with  clawed  digits;  carpus  and  tarsus 
cartilaginous;  endochondrium  well  formed. 

There  are  4  genera  which  constitute  this  family:  Urocordylus,  from  the  Coal 
Measures  of  Ireland;  Ceraterpeton,  from  the  Coal  Measures  of  Ireland  and  Eng- 
land; Diceratosaurus,  from  the  Coal  Measures  of  Linton,  Ohio;  Eoserpeton,  from 
the  Coal  Measures  of  Linton,  Ohio. 

These  may  be  distinguished  by  the  following  characters : 

I.  Skull  triangular,  truncated  behind,  with  rounded  muzzle  and  aborted  tabulare  cornua,  neural  spines  of 
caudal  vertebrae  long,  slender,  and  expanded  in  a  fan-like  manner;  tail  with  about  80  vertebra;; 
ventral  scutes  oat-like Urocordylus. 

II.  Skull  parabolic  and  of  great  width,  with  short  cornua  projecting  from  the  supratemporal ;  tabulare  cornua 
nearly  twice  as  long;  neural  spines  of  caudal  vertebrae  low  and  wide;  ventral  scutes  oblong;  caudal 
vertebras  about  50 Ceraterpeton. 

III.  Skull  broad  with  obtuse  snout,  tabulare  cornua  absent,  large,  pointed  posterior  expansions  from  supra- 

temporal,  posterior  table  within  the  cornua  truncate;  vertebrae  with  an  apical  sculptured  plate; 
caudal  vertebrae  numerous,  over  75;  ventral  scutellae  bristle-like,  arranged  en  chevron . .  Diceratosaurus- 

IV.  Skull  a  broad  oval,  with  large  posterior  projecting  supratemporal  horns,  posterior  table  of  skull  between 

cornua  truncate  without  the  small  lateral  projection  from  the  supratemporal,  orbits  a  long  oval, 
ribs  long,  curved  and  slender,  tail  unknown,  possibly  shorter  than  in  other  members  of  the 
family;  it  is  restored  as  short  in  Journal  of  Geology,  XVII,  p.  77,  fig.  20,  1909,  but  this  is  uncertain; 
the  skull  has  all  the  characters  of  the  family Eoserpeton. 

The  relationships  of  the  family  are  not  far  to  seek.  They  fall  in  immediately 
with  the  Amphibamidas  and  Hylonomida?  in  being  among  the  most  reptile-like  of 
the  Paleozoic  Amphibia.  The  group  is,  however,  distinctly  amphibian  in  the  pos- 
session of  4  fingers,  with  the  usual  microsaurian  phalangeal  formula. 

Genus  DICERATOSAURUS  Jaekel,  1903. 

Jaekel,  Ncucs  Jahrbuch  f.  Mineral.,  Geol.  u.  Palcon.,  Bd.  1,  p.  112,  1903. 
.Moodie,  Jour.  Geol.,  xvil,  pp.  63-69,  figs.  13-15,  1909. 

Type:  Diceratosaurus  punctolineatus  Jaekel. 

Orbits  in  the  anterior  two-thirds  of  the  axial  skull  length,  nostrils  near  to 
the  anterior  end  of  the  skull;  pineal  foramen  in  the  center  of  the  skull  roof;  skull 

115 


Il6  THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 

provided  with  tabulare  cornua  and  a  broad  backwardly  directed  process ;  quadrate 
angle  does  not  project  on  the  border  of  the  skull ;  sculpture  of  the  cranial  elements 
impressed  as  radial  grooves;  12  presacral  vertebrae,  1  sacral  with  expanded  neural 
spine  which  is  sculptured  at  the  top,  with  simple  long,  apparently  separately  ossi- 
fied transverse  processes ;  extremities  small ;  foot  with  5  digits ;  phalangeal  formula 
2-3-3-4-3- 

The  most  important  differences  between  Diceratosaurus  and  Ceraterpeton,  the 
most  nearly  allied  genus,  is  (in  Diceratosaurus)  in  the  more  anterior  position  and 
small  size  of  the  orbits,  the  backward  extension  of  the  quadrate  region,  and  the 
dorsal  expansion  of  the  vertebral  spines.  A  further,  and  more  important,  difference 
between  the  genera  is  in  the  location  of  the  backwardly  directed  processes  from  the 
skull.  In  Ceraterpeton  they  project  backward  from  and  are  a  portion  of  the  tabulare 
element,  while  in  Diceratosaurus  the  projection  consists  almost  entirely  of  squa- 
mosal and  supratemporal. 

Diceratosaurus  punctolineatus  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  1875,  p.  16. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  372,  pi.  xli,  fig.  4,  1875. 

Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  art.  xxv,  p.  356,  pi.  lxv,  1909. 

Type:  Specimen  No.  8606,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  was  first  described  by  Cope  as  Ceraterpeton  punctolineatum  (122). 
It  was  redescribed  on  more  abundant  material  (plate  19)  by  Jaekel  (347),  and  the 
following  is  taken  from  the  discussions  of  these  two  authors,  checked  by  my  own 
observations  on  the  type  specimen.  This  shows  a  portion  of  the  skull,  consisting 
of  the  squamosal  and  supratemporal  with  a  projecting,  convergent  horn.  The 
sculpturing  on  the  skull  is  similar  to  that  on  the  pectoral  plates,  of  which  there  are 
three  preserved  (plate  14,  fig.  4).  The  bones  of  the  fore  limbs  are  stout  and 
short.  The  ribs  are  only  slightly  curved.  The  character  of  the  vertebrae  can  not 
be  ascertained.  The  sculpturing  of  the  bones  consists  of  radiating  ridges,  grooves, 
and  pittings. 

Jaekel  (347)  described  from  the  museum  collection  at  Berlin  3  specimens  of 
this  species,  among  which  were  2  skulls.  There  were  associated  with  these  remains 
some  pectoral  plates  and  limb  bones  with  a  nearly  complete  series  of  vertebras.  A 
modified  translation  of  Jaekel's  description  follows: 

The  skull  of  the  largest  specimen  has,  including  the  horns,  a  length  of  35  mm. 
and  a  width  of  30  mm.  on  the  occipital  border.  From  the  anterior  end  to  the  pos- 
terior border  of  the  skull  (exclusive  of  the  horns)  there  is  a  length  of  25  mm.  The 
pineal  foramen  lies  about  midway  of  this  length.  The  orbits  are  rather  large,  al- 
most circular,  and  lie  about  midway  between  the  pineal  foramen  and  the  anterior 
border  of  the  skull.  The  nostrils,  which  lie  anterior  to  the  orbits,  are  a  rather  oblique 
oval  and  narrowed  on  the  lateral  ends.  The  distance  between  them  is  about  the 
same  as  that  between  the  orbits,  which  measures  7  mm. 

The  skull  roof  is  sculptured  with  pits  much  like  those  of  Archegosaurus.  The 
larger  the  bones,  the  rougher  the  sculpture.  The  bones  of  the  middle  of  the  skull 
(that  is,  the  parietals,  frontals,  premaxillae,  nasals,  and  postparietals)  are  of  the 


MOODIE 


PLATE  16 


I  •  Type  specimen  of  Diceratosaurus  punctolincatus  Cope,  from  Linton,  Ohio,  beds.  X  I .  Drawn  from  photo- 
graph. r/  =  claviclc;  A  =  humerus;  ic  =  in  terclavicle;  mc  =  metacarpals;  r-u  =  radius-ulna;  r6  =  rib; 
sc  =  scapula;  sq=  squamosal;  vs  =  ventral  scutella?. 

2.  Skull  of  Sauropleura  longidentata,  Moodie,  from  the  Coal  Measures  of  Linton,  Ohio.    X  I.    Drawn  from 

photograph,  fr  =  frontal ;  m  =  maxilla ;  n  =  nasal ;  or  -  orbit ;  par  =  parietal ;  pp  =  postparietal ;  tab  =  tabulare. 

3.  Mandible  of  Sauropleura  longidentata,  Moodie,  from  Coal  Measures  of  Linton,  Ohio.     X  i-5-     Drawn 

from  photograph. 

4.  Type  specimen  of  Sauropleura  enchodus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio.     X  1. 

5.  Additional  specimen  of  Diceratosaurus  punctolineatus  Cope,  from  the  Coal  Measures  of  Linton  Ohio. 

X  1.     '=  femur;  j'/  =  ilium;  mc  =  metacarpal;  mt  =  metatarsal;  pA  =  phalanx;  r  =  radius;  <  =  tibia. 


THE    MICROSAURIAN   FAMILY    UROCORDYLID^E.  117 

normal  form  and  only  show  an  unusual  difference  in  that  they  are  large  in  the 
inverse  order.  The  pineal  foramen  lies  in  the  anterior  third  of  the  parietals ;  that  is 
the  primitive  condition  which  occurs  in  the  young  forms  of  the  Branchiosauridae. 
The  nostrils  are  inclosed  by  the  premaxillae  in  front,  in  the  median  line  by  the  nasals 
and  laterally  by  the  maxillae.  The  jugals,  by  their  backward  prolongation,  form, 
behind  the  maxillae,  the  border  of  the  skull,  and  only  attain  to  some  size  on  the 
upper  side  of  the  cranium. 

Jaekel's  " perisquamosal "  (see  plate  15),  which  is  of  a  doubtful  nature,  is  not 
indicated  in  the  type  specimen  nor  in  the  specimens  of  the  other  two  species  (462) 
assigned  to  this  genus.  In  D.  robustus  and  D.  Icevis  the  "perisquamosal"  region  is 
easily  separable  into  its  component  elements.  The  sutures  between  the  elements 
may  have  been  indistinct  in  his  specimen,  but  it  is  hardly  conceivable  that  a  union 
of  the  skull  bones  would  occur  in  one  species  and  not  in  another  of  the  same  genus. 
Jaekel's  suggestion  (347)  that  "Etwas  mehr  Wahrscheinlichkeit  mochte  ich  der 
Vorstellung  beimessen,  dass  diese  Ausbreitungen  zum  Schutz  freier  Kiemen  dienten, 
wie  sie  z.  B.  bei  den  Perennibranchiaten  als  baumformige  Organe  weit  am  Halse 
herausragen"  can  hardly  find  acceptance  with  students  of  the  Paleozoic  Amphibia, 
since  there  is  not  the  slightest  evidence  that  the  Microsauria  ever  possessed  external 
gills  and  considerable  presumptive  evidence  that  they  did  not.  His  comparison 
of  the  "perisquamosal"  to  the  "Kiemendeckel"  of  the  fishes  is  also  very  unhappy 
on  morphological  grounds,  since  the  elements  of  his  "perisquamosal"  form  constit- 
uent parts  of  the  skull  roof,  which  the  operculum  never  does. 

The  palate  of  the  skull  (plate  15,  fig.  2)  has  been  determined  by  removing  the  skull 
bones  of  one  specimen.  Anteriorly  the  premaxillae  and  maxillae  are  clearly  recogniz- 
able as  large  dentiferous  elements.  The  premaxillae  have  4  to  7  teeth,  the  short  max- 
illa has  3  to  4.  All  the  teeth  are  of  nearly  equal  size.  Smaller  teeth  seem  to  be 
indicated  by  impressions  found  between  the  larger  ones.  The  vomers,  which  are 
tolerably  large,  unite  with  the  premaxillae  behind  and  inclose  at  least  half  of  the 
palatine  foramen  on  the  inner  side.  They  are  furnished  with  small  teeth,  which  in 
the  anterior  part  are  very  irregularly  placed,  but  they  are  more  regular  posteriorly. 

The  palatines  and  transverse  bones  are  questionably  identified.  They  seem  to 
lie  posterior  and  lateral  to  the  vomers,  but  the  sutures  are  indistinct.  The  large 
parasphenoid  seems  well  displayed  and  is  more  or  less  heart-shaped.  There  would 
seem  to  be  a  slight  indication  of  double  occipital  condyles.  The  pterygoids  are 
broad  plates  which  inclose  the  parasphenoid  and  form  the  lateral  boundary  of  the 
palate.    The  cotyli  are  very  indistinct,  but  appear  as  elongate  grooves. 

The  pectoral  girdle  (plate  15,  fig.  3)  consists,  apparently,  of  seven  elements,  three 
paired  and  one  unpaired.  The  unpaired  element  (the  interclavicle)  is  truncate  pos- 
teriorly and  acuminate  in  front,  with  its  surface  radially  grooved  and  the  anterior 
borders  beveled  for  articulation  with  the  clavicles.  The  clavicles  are  triangular,  as  is 
usual  with  the  Microsauria.  They  are  sculptured  with  radiate  grooves  and  ridges, 
with  decided  inosculations  at  the  ossific  center.  The  coracoids  have  only  part  of 
their  surface  ornamented ;  most  of  their  surface  is  smooth  for  articulation  with  the 
interclavicle  and  scapula.    A  long  spine  projects  from  the  inner  surface  of  the  cora- 


H8  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

coid.  A  pair  of  small  elements  lie  one  on  either  side  of  the  clavicles  and  Jaekel  (347) 
interprets  them  as  the  cleithra.  If  they  are  cleithra  they  are  unique  among  the 
Microsauria.  The  pectoral  girdle  does  not,  however,  indicate  that  Diceratosaarus 
is  "unique  among  all  known  quadrupeds." 

Jaekel  regards  the  limb  which  is  preserved  with  the  material  as  an  arm  (plate 
15,  fig.  4) ,  but  there  is  no  reason  stated  for  his  conclusion.  It  has  all  of  the  characters 
of  the  leg  and  may  be  regarded  as  such.  Only  a  part  of  the  lower  end  of  the  femur  is 
preserved.  The  tibia  and  fibula  are  preserved  as  separate  rod -like  elements  with  one 
of  the  bones  longer  and  larger,  probably  the  tibia.  There  are  5  toes  which  have  the 
customary  phalangeal  formula  for  a  microsaurian  foot  of  2-3~3-4(?)-3.  The 
tarsals  are  unossified. 

The  vertebrae  (plate  15)  were  perforated  for  the  notochord,  and  are  hour-glass- 
shaped,  with  the  neural  arch  thickened  to  support  a  heavy  spine  which  bore  a  sculp- 
tured plate.  These  apical  plates  occur  in  the  dorsal  region,  but  diminish  toward  the 
caudal  vertebrae.  The  number  of  the  vertebrae  in  the  dorsal  region  is  very  small  and 
in  the  tail  very  large.  There  are  possibly  2  vertebrae  in  the  cervical,  1 1  in  the  dorsal 
series;  the  thirteenth  carries  the  pelvis.    There  are  over  100  vertebrae  in  the  tail. 

The  ribs  have  an  expanded  head  and  the  transverse  processes  of  the  vertebrae 
are  long. 

The  following  account  is  based  on  the  writer's  study  of  the  type  specimen  and 
he  is  able  to  add  several  points  of  interest  to  a  knowledge  of  the  anatomy  of  the 
type  of  this  interesting  microsaurian. 

The  type  specimen  consists  of  1 1  consecutive  vertebrae  with  a  portion  of  the 
skull,  the  greater  portion  of  the  pectoral  girdle,  parts  of  both  fore  limbs,  ribs,  and 
ventral  scutellae  (plate  14,  fig.  4).  The  species  is  represented  in  the  collection  by  yet 
another  specimen,  on  which  Cope  based  his  Tuditamis  mordax  (plate  22,  fig.  5),  of 
which  he  himself  says :  ' '  Further  examination  of  the  specimen  on  which  the  latter 
(T.  mordax)  was  founded  leads  to  the  belief  that  it  is  an  imperfect  cranium  of  Cerater- 
peton  (Diceratosaurus)  punctolineatum  Cope."  The  plates  referred  to  are  rather  to  be 
regarded  as  elements  of  the  pectoral  girdle  and  I  believe  they  represent  the  clavicle 
and  a  portion  of  the  interclavicle. 

The  skull  of  the  present  species  is  fully  described  by  Jaekel.  The  type  specimen 
does  not  offer  any  evidence  in  support  of  Jaekel's  "perisquamosal,'.'  but  rather 
tends  to  the  idea  that  he  is  incorrect  in  his  assumption  of  the  fusion  of  these  elements 
of  the  skull.  The  direction  taken  by  the  ridges  and  grooves  on  the  elements  pre- 
served indicate  a  separation  between  the  supratemporal  and  the  squamosal.  I  do 
not  find  that  the  grooves  have  the  tendency  to  arise  from  a  common  center  of  ossi- 
fication in  the  squamosal,  as  suggested  in  the  figures  of  Jaekel.  The  horn  which 
projects  backward  from  the  squamosal  is  rather  large  and  heavy  for  the  size  of  the 
skull,  and  after  curving  slightly  inward  ends  in  a  blunt  point  and  not  sharply,  as 
Jaekel  figures  in  his  specimens.  The  vertebral  column  is  indistinctly  preserved  and 
I  have  nothing  to  add  to  Jaekel's  account  given  above. 

In  the  structure  of  the  pectoral  girdle  my  results  are  greatly  at  variance  with 
those  of  Jaekel.     I  do  not  find  the  remarkable  elements  which  Jaekel  has  figured 


THE    MICROSAURIAN    FAMILY    UROCORDYLID/E. 


119 


(347)  in  his  specimens.  On  the  other  hand,  I  find  a  normal  microsaurian  pectoral 
arch  (464),  such  as  has  been  described  for  numerous  other  forms.  There  are  present, 
distinctly  preserved  in  the  type  specimen,  the  scapulae,  the  clavicles,  and  the  interclav- 
icle,  with  the  possibility  of  the  coracoid.  The  peculiar  element  referred  to  by  Cope 
as  resembling  a  "lacertilian  pubis"  is  without  doubt  the  left  scapula  of  the  animal 
(plate  16,  fig.  1 ) .  Its  form  compares  very  favorably  with  that  of  Ceraterpeton  as  fig- 
ured by  Woodward  (630).  The  coracoid  may  be  represented  by  the  fragment  which 
lies  close  to  the  scapula.  The  sculptured  element  lying  next  to  the  supratemporal 
horn  of  the  skull  is  the  right  clavicle  preserved  bottom  side  up.  Of  the  other  two 
sculptured  elements,  one  is  the  interclavicle,  only  a  portion  of  which  is  preserved. 
The  left  clavicle  lies  beside  it.  The  clavicles  in  this  species  have  a  tendency  to 
assume  the  triangular  shape  so  common  in  other  species  of  Microsauria,  and  the 


Fig.  24. 

A.  Skull  of  Diceratosaurus  lozvis  Moodie,  from  the  Linton  Coal  Measures.  X  I.  /.frontal: 
j,  jugal;  mx,  maxilla;  n,  nasal;  or,  orbit;  par,  parietal;  pof,  postfrontal;  po,  postorbital; 
pf,  prefrontal;  pp,  postparietal ;  sq,  squamosal;  spt,  supratemporal;  qj,  quadratojugal ; 
pmx,  premaxilla;  tab,  tabulare. 

B.  Reconstruction  of  skull  outlines  of   Diceratosaurus  robuslus  Moodie,   from  the  Coal 

Measures  of  Ohio.  X  0.75.  fr,  frontal;  7,  jugal;  or,  orbit;  par,  parietal;  pof,  postfrontal; 
po,  postorbital;  pp,  postparietal;  qj,  quadratojugal;  spt,  supratemporal;  tab,  tabulare. 

interclavicle,  so  far  as  can  be  determined,  was  shield-shaped.  The  upper  sur- 
faces of  the  pectoral  elements  are  marked  by  grooves  for  the  attachment  of  the 
pectoral  muscles. 

The  ventral  scutellation  is  present  in  a  small  patch  (plate  16,  fig.  i)  near  the 
horn  of  the  skull.  The  scutse  are  oat-shaped  and  take  the  usual  form.  The  ribs 
are  not  long,  are  rather  stout,  and  beyond  the  proximal  curve  are  nearly  straight  to 
the  obtuse  tips.  The  heads  of  the  ribs  are  so  obscure  that  it  is  impossible  to  deter- 
mine whether  they  were  two-headed  or  not.  They  are  expanded  proximally  and 
there  is  a  slight  tendency  to  a  division  of  the  head. 

Portions  of  both  fore  limbs  are  preserved.  The  right  limb  possesses  the  hume- 
rus, separate  radius  and  ulna,  and  2  metacarpals.  The  other  possesses  only  the 
radius,  3  metacarpals,  and  a  portion  of  a  phalanx.  The  humerus  is  a  very  stout 
bone  and  at  once  recalls  that  of  Amblyrhynchus.    The  ends  are  expanded  and  there 


120  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

are  roughnesses  on  the  bone  for  the  attachment  of  muscles.  The  radius  and  ulna 
are  subequal  in  size.  They  are  both  expanded  more  proximally  than  distally.  The 
carpus  was  cartilaginous.  An  additional  specimen  of  this  species  is  figured  on  plate 
1 6,  fig.  5.    This  adds  to  our  knowledge  of  the  pelvis  especially. 

Measurements  of  the  Type. 

mm.  mm. 

Length  of  entire  specimen 80  Width  of  various  portions  of  ulna  same  as  radius. 

Length  of  tabulare  horn  of  skull 20         Length  of  the  only  phalanx  preserved 5 

Width  at  base 4         Length  of  vertebra 5 

Width  at  tip 2.5      Width  of  vertebra 4 

Length  of  right  humerus 16         Length  of  longest  rib 17 

Width  at  middle  of  shaft 3         Width  of  rib  at  widest  part 1.5 

Width  at  proximal  end 5         Width  of  clavicle 18 

Width  at  distal  end 5.5      Length  of  clavicle 20 

Length  of  radius 9  Length  of  interclavicle 25 

Length  of  ulna 9         Width  of  interclavicle 16 

Width  of  radius  at  proximal  end 2.5      Length  of  single  side  of  chevron  scute 7 

Width  at  middle 1.5  Width  of  same 25 

Width  at  distal  end 2 

The  specimen  (No.  2566,  Am.  Mus.  Nat.  Hist.)  on  which  Cope  based  his 
Tuditanus  mordax  is  composed  of  two  plates  of  the  above-described  species. 

Diceratosaurus  laevis  Moodie. 
Moodie,  Jour.  Geol.,  xvn,  No.  I,  p.  63,  figs.  13,  14,  1909. 

Type:  Specimen  No.  102  (8680  G),  American  Museum  of  Natural  History, 
where  it  forms  part  of  the  Newberry  collection. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  is  represented  by  an  almost  complete  skull,  which  had  been  identi- 
fied previously  by  Cope  as  Tuditanus  radiatus.  The  specimen  consists  of  the 
impressions  of  the  bones  of  the  cranial  roof,  the  bones  themselves  having  disap- 
peared. It  is  not  probable  that  the  dorsum  of  the  skull  was  smooth.  The  details 
in  the  structure  of  the  skull  have  been  ascertained  quite  definitely.  There  can  be 
no  doubt  that  the  arrangement  of  the  elements  is  accurate,  as  shown  in  figtire  24  A. 
The  supratemporal,  as  in  Erpetosaurus  tabulatus  Cope,  is  excluded  from  the  parietal. 

The  form  of  the  skull  at  once  recalls  that  of  the  species  D.  punctolineatus,  as 
figured  by  Jaekel  (see  plate  1 5) .  The  orbits  are  located  in  nearly  the  same  region 
of  the  skull  and  the  sutures  separating  the  cranial  elements  are  quite  similar  in  the 
anterior  portions.  The  species  D.  loms  is  based  on  the  divergent  character  of  the 
horn-like  protuberances  which  project  from  the  squamosals.  The  horns  of  D, 
punctolineatus  are  convergent.  The  present  skull  is  also  smaller  and  the  parietals 
in  D.  leans  are  much  larger  than  in  the  type  species.  In  the  type  species,  also,  the 
pineal  foramen  is  located  well  forward  in  the  parietal,  while  in  the  present  form  the 
foramen  is  located  well  posterior. 

The  skull  is  almost  rectangular.  The  nostrils  are  elongate  ovals.  The  orbits 
are  circular  and  the  distance  between  them  is  equal  to  two-thirds  of  the  dimensions 
of  the  orbit.  They  are  located  well  forward  in  the  skull  and  are  bounded  laterally 
by  the  maxillaries.  The  nostrils  have  much  the  same  character  as  in  the  type  form, 
being  broadly  oval. 

The  premaxillae  are  elongate  transversely,  being  about  twice  as  long  as  wide. 
They  are  identical  in  shape  and  relations  with  the  same  elements  in  D.  punctoline- 
atus Cope.  The  nasal  is  nearly  square  and  forms  the  interior  boundary  of  the  nos- 
tril.   The  frontal  is  elongate  in  the  median  length  of  the  skull  and  it  is  acuminate 


THE   MICROSAURIAN   FAMILY   UROCORDYLIOE.  121 

posteriorly,  where  the  acumination  is  inclosed  by  the  parietal  and  postfrontal.  The 
parietals  are  by  far  the  largest  elements  in  the  cranium.  They  form  together  an 
oval  which  is  elongate  in  the  longitudinal  diameter  of  the  skull.  They  inclose 
between  them,  in  the  median  suture,  the  small  pineal  foramen.  They  are  acu- 
minate in  front,  with  a  broad  truncate  posterior  base,  where  they  are  bounded  by  the 
postparietals.  The  postparietal  is  nearly  square,  being  somewhat  wider  than  long. 
It  joins  the  tabulare  and  the  parietal.  The  tabulare  is  elongate  in  the  long  diame- 
ter of  the  skull.  It  ends  anteriorly  in  a  point  which  is  inserted  between  the  post- 
orbital  and  the  parietal,  and  bears  a  short  protuberance  posteriorly,  much  as  does 
the  same  element  in  the  type  species. 

There  are  four  elements  which  take  part  in  the  formation  of  the  posterior  border 
of  the  skull.  These  are  the  postparietal,  the  tabulare,  the  squamosal,  and  the 
supratemporal.  It  is  very  unusual  for  the  supratemporal  to  reach  the  posterior  edge 
of  the  cranium.  The  prefrontal  lies  anterior  to  the  orbit,  of  which  it  forms  the  ante- 
rior border.  The  lacrimal  has  not  been  detected,  although  Jaekel  (347)  has  indi- 
cated it  in  his  drawings  of  the  skull  of  the  type  species.  The  maxilla  is  elongate  and 
forms  the  lateral  border  of  the  skull.  No  teeth  have  been  detected,  although  they 
were  doubtless  the  same  as  Jaekel  has  figured  in  D.  punctolineatus.  The  jugal  is  an 
elongate  element  joining  the  maxilla  posteriorly.  Jaekel  included  this  element  in 
his  "perisquamosal,"  but  the  sutures  are  clearly  evident  in  the  present  specimen 
and  there  is  no  evidence  of  a  structure  at  all  similar  to  a  "perisquamosal."  The 
postorbital  is  fully  as  large  as  the  jugal  which  it  joins,  forming  a  part  of  the  poste- 
rior border  of  the  orbit  and  ending  posteriorly  in  a  point  which  is  inclosed  by  the 
tabulare  and  the  squamosal.  The  postfrontal  with  the  foregoing  element  forms 
the  entire  posterior  border  of  the  orbit  and  it  likewise  ends  in  a  point  inclosed  by 
the  parietal  and  the  postorbital.  The  quadratojugal  has  much  the  same  shape  and 
relations  as  in  D.  punctolineatus,  although  it  is  located  further  back.  The  squa- 
mosal is  also  elongate,  as  are  most  of  the  posterior  cranial  elements,  and  it  also 
has  an  acumination  which  is  directed  forward  and  is  inclosed  by  the  postorbital 
and  jugal.  The  anterior  suture  of  this  element  is  rather  indistinct,  but  it  is,  I 
believe,  as  represented  (fig.  24).  The  element  is  elongate  and  is  prolonged 
posteriorly  to  form  the  horn,  which  ends  in  a  blunt  point  and  is  not  sharp,  as  in 
the  type  species. 

Jaekel  (347)  regards  the  species  Dicer  atosaur  us  punctolineatus  Cope  as  being 
unparalleled  among  known  vertebrates  in  the  possession  of  a  "perisquamosal"  ele- 
ment. In  closely  allied  species  the  "perisquamosal"  is  easily  separated  into  its 
component  elements,  and  the  morphology  of  the  present  skull  would  throw  con- 
siderable doubt  on  Jaekel's  interpretation  of  the  skull  of  the  type  species.  Another 
specimen,  described  below  as  another  species  of  this  genus,  shows  no  evidence  of 
this  fusion.  So  far  as  I  can  learn,  there  have  been  no  cases  of  true  fusion  of  cra- 
nial elements  correctly  reported,  unless  it  be  that  which  possibly  exists  between 
the  two  frontals  in  the  skull  of  Diplocaulus.  It  was  on  the  basis  of  such  fusions 
that  Maggi  (397)  proposed  to  derive  the  interparietals  of  the  primates  from  the 
tabulare  of  the  stegocephalians. 


122  THE    COAL   MEASURES    AMPHIBIA   OF   NORTH   AMERICA. 

The  posterior  outline  of  the  skull  in  the  present  specimen  is  not  well  preserved 
and  the  outline  as  given  may  be  slightly  inaccurate.  The  indentation  figured  by 
Jaekel  in  the  posterior  border  of  the  skull  of  the  type  form  is  not  present  in  the 
species  under  discussion. 

Measurements  of  the  Type  Skull  of  Diceratosaurus  l^evis  Moodie. 

ram.  mm. 

Length  of  skull  along  median  suture 37  Width  of  skull  across  the  orbits 30 

Length  from  muzzle  to  tip  of  horn 50  Interorbital  width 6 

Width  between  tips  of  horns,  estimated 40  Length  of  nostril  opening 2 

Width  of  orbit 7  Width  of  nostril 1 

Length  of  orbit 10  Diameter  of  the  pineal  foramen —   I 

Diceratosaurus  robustus  Moodie. 

Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  67,  fig.  15,  1909. 

Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  art.  xxv,  p.  355,  pi.  lxiii,  fig.  2,  1909. 

Type:  Specimen  No.  8611  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  present  species  is  indicated  by  the  left  portion  of  a  cranium  representing  a 
large  individual.  The  characters  of  the  skull  are  so  clearly  marked  that  it  seems 
worthy  of  description.  The  presence  of  horns  as  given  in  the  restoration  of  the 
skull  (fig.  24,  B)  is  based  on  the  analogy  with  the  other  two  species  of  this  genus,  in 
both  of  which  horns  are  present.  The  generic  determination  of  the  species  is  based 
on  the  large  size  of  the  postorbital,  which  is  essentially  characteristic  of  the  other 
species  of  Diceratosaurus. 

The  characters  which  distinguish  the  species  from  others  of  the  genus  are  the 
large  postorbitals  and  the  small  parietals,  which  are  excluded  from  union  with  the 
postf rentals  on  account  of  the  large  size  of  the  frontal.  In  the  other  two  known 
species  the  frontal  is  small  and  the  parietal  comes  forward  to  join  the  post  frontal. 
The  present  species  exhibits  a  skull  which  is  nearly  twice  as  large  as  that  of  D. 
IcBvis  and  nearly  three  times  the  size  of  the  skull  of  D.  punctolineatus. 

The  portion  of  the  skull  preserved  shows  the  cranium  to  have  had  a  rather  acu- 
minate snout,  not  blunt  as  in  the  type  species.  The  orbit  is  an  elongate  oval, 
although  it  has  the  same  relative  position  in  the  skull  as  in  the  other  species.  The 
nostril  is  indicated  by  an  oval  depression  near  the  anterior  edge  of  the  skull.  The 
frontals,  as  indicated  by  the  sutures  present  on  the  portion  of  the  skull  which  is 
preserved,  are  fully  as  long  as  the  parietals.  Whether  they  were  as  wide  as  is  repre- 
sented is  uncertain.  The  postfrontals  are  very  small  bones,  the  sutures  of  which 
are  somewhat  uncertain,  although  they  can  not  be  far  from  what  is  represented 
(fig.  24,  B) .  The  postorbital  is  large  and  elongate,  and  is  distinctive  of  this  species  on 
account  of  its  unusual  size,  although  it  does  not  attain  the  same  proportions  as  in 
other  members  of  the  genus.  The  parietals  are  elongate  and  narrow.  The  pineal 
foramen  is  represented  by  its  lateral  edge  and  its  position  is  about  midway  of  the 
longitudinal  diameter  of  the  parietals.  The  narrow  postparietal  is  represented  by 
its  anterior  border;  as  restored  (fig.  24,  B)  it  may  be  too  long.  The  tabulare,  also,  is 
represented  by  an  anterior  portion  and  it  shows  this  element  to  have  the  position 
and  form  which  is  typical  of  the  form  Diceratosaurus  Icevis.  Such  other  of  the 
cranial  elements  as  are  indicated  are  based  on  the  relations  discovered  in  D.  Iceins. 


THE   MICROSAURIAN   FAMILY   UROCORDYLIOE.  1 23 

The  heavy  line  on  the  left  of  the  drawing  (fig.  24,  B)  represents  the  outline  of  the 
preserved  portion.  The  skull,  as  restored,  may  be  a  little  too  long,  and  the  shape  of 
the  horns  is  conjectural.  In  the  orbit  there  are  preserved  2  misplaced  teeth  show- 
ing longitudinal  fluting.    The  longest  tooth  is  about  3  mm. 

Measurements  of  the  Type  Skull  of  Diceratosaurus  robustus  Moodie. 

mm.  mm. 

Median  length  of  skull,  estimated 67      Length  of  longest  tooth 3 

Posterior  width  of  skull,  estimated 78      Width  of  same  tooth  at  base 1.5 

Length  of  orbit 18      Length  of  postorbital 27 

Width  of  orbit 12      Width  of  postorbital 14 

Genus  EOSERPETON  Moodie,  1909. 

Moodie,  Jour.  Geol.,  xvn,  pp.  76-79,  fig.  20,  1909. 

Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  p.  355,  pi.  lxiii,  fig.  1,  1909. 

Type:  Eoserpeton  tenuicorne  Cope. 

The  genus  was  proposed  for  the  reception  of  a  single  species  originally  referred 
by  Cope  to  Ceraterpeton  (C.  tenuicorne).  The  species  can  not  be  placed  in  the  genus 
Ceraterpeton  on  account  of  the  form  and  structure  of  the  skull,  which  varies  widely 
from  that  of  the  type  species,  Ceraterpeton  galvani  Huxley,  from  the  Kilkenny  Coal 
Measures  of  Ireland.  The  most  important  character  in  which  the  present  species 
differs  from  C.  galvani  is  the  peculiar  form  taken  by  the  squamosal  and  by  the  posi- 
tion of  the  "horn."  These  characters  will  be  evident  on  referring  to  figure  25.  No 
undoubted  remains  of  Ceraterpeton  have  been  found  outside  the  British  Isles.  Fritsch 
referred  (251)  a  species,  previously  described  as  Scincosaurus  crassus,  to  this  genus, 
but  Andrews  (8),  Jaekel  (347)  and  Woodward  (630)  all  unite  in  placing  the  species 
in  the  genus  where  it  was  formerly  described.  Jaekel  even  says  that  the  Scinco- 
saurus has  no  horns,  so  far  as  he  can  determine.  Cope  referred  3  species  (123) 
from  the  Linton  Coal  Measures  of  Ohio  to  the  genus  Ceraterpeton,  but  it  has  been 
shown  elsewhere  (347)  that  no  one  of  them  belongs  in  the  genus,  nor  in  fact  do 
they  all  belong  in  one  genus. 

Eoserpeton  tenuicorne  Cope. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  pp.  372-373,  pi.  xlii,  fig.  2,  1875. 
Cope,  Proc.  Am.  Phil.  Soc,  xxn,  p.  407,  1885. 
Cope,  Proc.  Am.  Phil.  Soc,  xxxvi,  p.  85,  pi.  iii,  fig.  2,  1897. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  vol.  37,  p.  23,  1909. 

Type :  Specimen  in  the  American  Museum  of  Natural  History.  There  are  also 
specimens  Nos.  4472  and  4473  in  the  U.  S.  National  Museum. 

Locality  and  horizon:  Linton,  Ohio,  Coal  Measures. 

The  species  was  founded  on  a  complete  skull  preserved  on  obverse  sides  of  a 
block  of  coal.  Cope  (123,  pi.  xlii,  fig.  2)  figured  this  skull.  The  figure  is  poorly 
executed  and  does  not  do  justice  to  the  specimen,  which  is  really  well  preserved. 
In  general  the  skull  is  oval,  with  the  orbits  located  well  towards  the  acuminate 
snout.  The  interorbital  space  is  equal  to  twice  the  width  of  the  orbit.  The  pineal 
foramen  lies  near  the  center  of  the  skull.  The  quadrate  angles  are  drawn  out  into 
slender  acuminate,  longitudinally  striate  horns,  processes  from  the  squamosal. 
The  "horn"  arises  from  an  expanded  base,  which  is  a  portion  of  the  cranial  element 
at  the  postero-lateral  angle  of  the  skull.    This  character  is  taken  as  the  distinctive 


124 


THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


one  of  the  genus.    It  is  possessed  by  no  other  form  of  Carboniferous  air-breathing 
vertebrate,  in  association  with  the  other  characters  of  the  form. 

The  boundaries  of  the  premaxillse  are  indefinite,  but  what  remains  of  the  sutures 
indicates  that  the  elements  were  small.  No  teeth  have  been  detected.  The  nasal 
is  likewise  not  clearly  defined,  but  the  frontal  is  an  elongate  element  which  occu- 
pies the  space  between  the  orbits  and  joins  the  parietal  posteriorly.    The  parie- 


Measurements  of  the  Tyj>e  of  Eoserfe- 
ton  tenuicorne. 

mm. 

Median  length  of  skull 26 

Width  across  squamosal  enlargement..  .  30 

Width  across  base  of  horns 25 

Length  from  muzzle  to  tip  of  horn 40 

Length  of  horn 10 

Width  of  horn  at  base 4 

Length  of  orbit 4 

Width  of  orbit 3 

Interorbital  space 6 


Nos.  4472  and  4473,  U.  S.  National  Museum, 
from  Linton,  Ohio,  Coal  Measures. 

mm. 

Median  length  of  skull 16 

Maximum  width  of  skull 20 

Length  of  horn  from  base 7 

Length  of  orbit 3 

Interorbital  width 3.5 

Length  of  vertebral  column  to  sacrum . .  33 

Length  of  femur 5.5 

Length  of  tibia  and  fibula 3 

Length  of  second  digit  (incomplete) 7 

Length  of  metatarsal 1 .5 

Length  of  clavicle 6 

Width  of  clavicle 3.5 


6   tff 

I    C      1 

/ 


w 

H 


Fig.  25. — Restoration  of  skeleton  of  Eoserpeton  lenuicorne  Cope.     X  1 . 5. 


Skull:  pmx,  premaxilla;  n,  nasal;  fr,  frontal;  par,  parietal;  pf,  prefrontal;  nut,  maxilla;  pof,  postfrontal;  po,  post- 
orbital;  pp,  postparietal;  j,  jugal;  qj,  quadratojugal;  spt,  supratemporal ;  sq,  squamosal;  tab,  tabulare. 

Skeleton:  ic,  interclavicle;  cl,  clavicle;  sc,  scapula;  h,  humerus;  r,  radius;  u,  ulna;  c,  carpus;  sr,  sacral  rib  (un- 
certain?); il,  ilium;  /,  femur;  fb,  fibula;  /,  tibia;  ts,  tarsus. 

tals  form  a  large  oval  space,  so  characteristic  of  many  of  the  Carboniferous  Micro- 
sauria,  in  the  anterior  third  of  which  occurs  the  median  parietal  foramen.  The 
postparietal  is  almost  square,  and  forms  part  of  the  posterior  boundary  of  the  skull. 
The  tabulare  has  the  usual  position  and  relations.  The  prefrontal  is  ill  defined. 
The  postfrontal  is  small  and  forms  a  slender  rod  on  the  postero-inner  boundary  of 
the  orbit.    The  postorbital  is  small  and  its  bounding  suture  with  the  postfrontal  is 


THE   MICROSAURIAN   FAMILY   UROCORDYLIOE.  125 

indefinite.  The  jugal  is  only  partially  represented  in  the  specimen,  and  that  part 
forms  the  outer  boundary  of  the  orbit.  The  maxillary  sutures  are  not  defined. 
There  are  no  evidences  of  teeth,  since  the  skull  is  compressed  dorso-ventrally.  The 
quadratojugal  is,  apparently,  a  larger  element  than  usual,  with  the  usual  rela- 
tions. The  supratemporal  lies  in  great  part  in  front  of  the  squamosal,  but  still  has 
the  normal  relations  of  that  element.  The  squamosal  is  the  characteristic  feature 
of  the  skull.  It  is  very  tumid  at  its  base  and  projects  into  a  long,  slender,  acumi- 
nate horn,  the  tumid  portion  being  ornamented  by  radiating  striae. 

Another  specimen  of  this  species  presents  the  greater  part  of  the  skeleton.  How- 
ever, very  little  can  be  added  to  our  knowledge  of  the  skull  structure.  It  is  barely 
possible  that  the  second  specimen  may  be  distinct  from  the  type.  The  horns  are 
curved  inward,  but  otherwise  there  is  little  or  no  difference.  One  of  the  most  inter- 
esting and  important  features  of  the  complete  specimen  is  the  unusual  preservation 
of  a  leg,  with  impressions  of  15  or  more  vertebras,  and  traces  of  curved  ribs  which 
are  intercentral  in  position. 

The  femur  is  slender  and  expanded  at  the  ends,  with  the  articular  surfaces  well 
formed.  The  tibia  and  fibula  are  mere  rods  of  bone,  although  the  tibia  has  slightly 
expanded  extremities.  There  is  no  osseous  tarsus.  There  are  5  digits  in  the  foot; 
the  second  one  is  entire  and  contains  4  phalanges;  the  other  digits  are  incomplete. 
The  foot  is  remarkably  long  and  slender,  and  is  fully  as  long  as  the  tarsal  space  plus 
the  tibia,  with  the  terminal  phalanx  clawed. 

There  are  impressions  of  2  oval  and  elongate  clavicles  in  the  pectoral  region. 
The  outer  end  is  not  expanded  as  is  usual,  and  the  surface  is  ornamented  with 
grooves  and  ridges  which  radiate  from  a  common  center. 

The  entire  remains  measure  scarcely  3  inches  in  length  and  it  is  to  be  doubted 
if  the  creature  attained  a  length  of  more  than  4  inches.  It  is  probably  a  young 
form,  but  there  are  no  evidences  of  external  gills.  The  chevron  armature  is  but 
poorly  preserved,  but  so  far  as  can  be  determined  it  is  not  different  from  that  of 
other  Microsauria,  such  as  Amphibamus. 


CHAPTER  XVII. 

THE    M1CROSAUR1AN    FAMILY    AMPH1BAMID/E,    FROM    THE    COAL    MEASURES 

OF  MAZON  CREEK,  ILLINOIS. 

Family  AMPHIBAMID^  new  family. 

Small,  lizard-like,  terrestrial  or  semi-aquatic,  megacephalic  microsaurians, 
known  from  3  species.  The  family  characters  are  the  huge  size  of  the  head  as 
compared  to  the  body,  the  short,  stumpy  body  with  about  25  short  dorsal  verte- 
bras, a  very  short  tail,  phalanges  clawed,  pubis  of  calcified  cartilage,  sclerotic 
plates  in  the  orbit  to  the  number  of  29  or  30  in  each,  ventral  armature  well 
developed.     Teeth  anisodont,  sharp,  conical,  non-striate. 

Two  genera  are  associated  in  this  family:  Amphibamus  grandiceps  Cope,  known 
from  three  nearly  complete  skeletons;  A.  thoracatus  Moodie,  known  from  a  single 
incomplete  skeleton;  Cephalerpeton  ventriarmatum  Moodie,  anterior  portion  of  body 
and  skull.  The  species  are  all  from  the  Mazon  Creek  shales  and  the  family  seems 
unrepresented  elsewhere.  It  may  be  necessary  to  compare  the  Amphibamidae  with 
the  Hylonomidas  when  the  latter  group  is  better  known,  but  in  the  light  of  our 
present  knowledge  the  two  families  are  distinct. 

The  genera  may  be  distinguished  as  follows: 

I.  Size  small,  less  than  3  inches  in  total  length,  skull  with  deeply  incised  tympanic  notches  (ear-slits)  .  .Amphibamus 
II.  Size  relatively  large,  body-length  6  inches  or  more,  teeth  distinctly  anisodont,  skull  with  nearly  even 

posterior  table,  limbs  very  long,  ventral  armature  highly  developed Cephalerpeton 

Genus  AMPHIBAMUS  Cope,  1865. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  1865,  pp.  134-137.    Geol.  Surv.  Ills.,  11,  pp.  135-141,  pi.  xxxii,  1  text-fig. 
Hay,  Proc.  Am.  Phil.  Soc,  xxxix,  p.  120,  1900. 
Moodie,  Jour.  Geol.,  xvn,  p.  81,  fig.  24,  1909. 

Type:  Amphibamus  grandiceps  Cope. 

The  publication  of  the  type  species  of  this  genus  began  the  researches  of  Pro- 
fessor Cope  on  the  extinct  Amphibia  of  North  America,  which  he  continued  for  so 
many  years  with  such  excellent  results  (105-177).  The  description  was  based  on  a 
single  specimen  (plate  3,  fig.  7)  belonging  to  Mr.  Joseph  Evans,  of  Morris,  Illinois, 
who  loaned  it  to  Dr.  Worthen  for  the  Illinois  Geological  State  Survey  (107),  in  order 
that  it  might  be  described.  The  type  has  been  destroyed  by  fire;  so  I  am  informed 
by  Mr.  L.  E.  Daniels,  of  Rolling  Prairie,  Indiana.  There  are  two  other  known  speci- 
mens of  the  species.  One  is  in  the  collection  of  Mr.  Daniels  and  the  other  No.  794, 
of  Yale  University  Museum. 

This  genus  may  be  clearly  separated  from  all  the  other  microsaurians  by  char- 
acters which  are  peculiar  to  the  form.  Among  these  may  be  mentioned  the  posses- 
sion of  sclerotic  plates  in  the  eyes;  the  large  size  of  the  orbits  in  comparison  with 
the  dimensions  of  the  skull;  the  short,  broad  form  of  the  body;  the  very  short  tail; 
the  possession  of  a  calcified  cartilaginous  pubis;  clawed  phalanges;  presacrals  22. 
The  character  which  places  the  genus  distinctly  in  the  Microsauria  is  the  possession 
of  long,  slender,  curved  ribs,  first  detected  on  Mr.  Daniels's  specimen  (plate  14,  figs. 

126 


MOODIE 


PLATE   17 


Type  of  Saurerpeton  latithorax  Cope.      X  1.5.     Original  in  U.  S.  National  Museum. 


THE    MICROSAURIAN    FAMILY   AMPHIBAMIOE.  127 

1,2),  by  Dr.  Hay  (316).  Its  stegocephalian  characters  are  evident  in  every  partic- 
ular of  its  anatomy — the  roofed  skull,  the  arrangement  of  the  cranial  elements,  the 
presence  of  a  well-developed  ventral  armature,  and  the  digital  formula  (4  for  the 
hand  and  5  for  the  foot). 

The  genus  A  mph ibamus  was  regarded  by  Cope  as  a  representative  of  a  new  order 
of  vertebrates  which  he  called  (105)  Xenorachia.  He  later  ( 1 23)  abandoned  this,  how- 
ever. Fritsch  (251),  Zittel  (642),  and  others  regarded  Amphibamus  as  a  branchio- 
saurian.  The  exact  position  of  the  form  was  uncertain  until  1900,  when  Dr.  Hay  (316) 
described  the  long,  curved  ribs  and  suggested  its  place  among  the  Microsauria.  He, 
however  (Cat.  Foss.  Vert.,  p.  410),  made  the  mistake  of  including  the  branchio- 
saurian  family  Protritonidae,  under  Microsauria,  thus  confusing  the  subject  further. 
The  genus  (462)  has  not  the  slightest  relationship  with  the  Branchiosauria. 

Amphibamus  grandiceps  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  pp.  134-137,  1865;  Geol.  Surv.  Ills.,  11,  pp.  135-141,  pi.  xxxii,  and  1  wood- 
cut, 1866. 

Hay,  Proc.  Am.  Phil.  Soc,  xxxix,  p.  120,  1900. 

Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  82,  fig.  24,  1909. 

Moodie,  Kan.  Univ.  Sci.  Bull.,  vi,  No.  2,  pp.  343-349,  pi.  1,  figs.  1  and  2;  pi.  5,  fig.  3;  pi.  7,  fig.  1;  pi.  n, 
12,  13,  1912. 

Type:  Specimen  has  been  destroyed.  There  is  an  excellent  specimen  (plate  4, 
figs.  5,  6),  No.  794  (1234),  in  Yale  University  Museum,  and  another  nearly  as  good 
in  the  possession  of  Mr.  L.  E.  Daniels,  of  Rolling  Prairie,  Indiana. 

Horizon  and  locality:  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  form  of  the  skull  of  Amphibamus  grandiceps  Cope  is  not  unlike  that  of 
Tuditanus  minimus  Moodie  (462)  from  the  Linton,  Ohio,  beds,  but  it  is  less  acumi- 
nate than  in  that  form.  The  large  size  of  the  orbits  is  especially  striking.  The  shape 
of  the  skull  is  triangular,  with  concavities  in  the  posterior  table  which  correspond 
to  the  ear-slits  so  characteristic  of  Metoposaurus  (242)  from  the  Keuper  of  Germany. 
The  narrowed  posterior  table  of  the  skull  is  truncate,  as  in  several  other  genera  of 
Microsauria,  notably  Tuditanus  and  Saurerpeton.  In  structure  the  skull  differs 
but  little  from  many  of  the  other  Carboniferous  forms,  but  the  arrangement  of  the 
elements  of  the  skull  is  more  regular  than  in  other  genera. 

The  premaxillaries  are  very  small  elements  in  the  anterior  tip  of  the  skull.  They 
border  the  nares.  The  skull  is  rather  peculiar  among  the  Microsauria  in  the  pos- 
session of  a  distinct  lacrimal.  I  have  detected  this  element  in  the  cranium  of 
Stegops  divaricata  Cope.  As  here  defined  the  lacrimal  is  triangular,  with  its  pos- 
terior border  formed  exclusively  by  the  prefrontal.  Its  other  relations  are  the  nor- 
mal ones.  The  nasal  is  elongate,  with  the  usual  relations  of  that  element.  The 
frontal  is  slightly  longer  and  broader  than  the  nasal.  It  apparently  forms  a  por- 
tion of  the  inner  border  of  the  orbit.  The  parietal  foramen  lies  in  the  anterior 
fourth  of  the  parietal,  a  rather  unusual  position  for  this  structure.  The  parietals, 
as  in  so  many  of  the  Microsauria,  together  form  the  largest  element  of  the  skull  and 
are  roughly  a  triangular  area  in  the  postero-median  portion  of  the  skull.  The  post- 
parietal  and  the  tabulare  are  clearly  distinguishable  and  they  have  the  usual  rela- 
tions for  those  elements.  The  maxillary,  jugal,  and  quadratojugal  together  form 
the  greater  part  of  the  maxillary  border.    The  postero-lateral  angle  of  the  skull  is, 


128 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


as  usual,  formed  by  the  squamosal.  The  orbit  is  bounded  posteriorly  by  the  post- 
orbital  and  the  postfrontal,  which  include  in  the  angle  between  them  the  quad- 
rangular squamosal.  The  orbit 
is  especially  remarkable  for  its 
size  as  compared  with  the  dimen- 
sions of  the  skull,  being  without 
a  parallel  among  other  known 
Microsauria.  Around  the  border 
of  the  orbit  in  the  specimen  Cope 
studied  (105)  there  were  found 
14  quadrangular  plates  which 
he  called  "superciliary  plates." 
Hay  (316)  was  inclined  to  regard 
them  as  sclerotic  plates.  In  the 
Yale  Museum  specimen  (plate  4, 
figs.  5,  6)  there  are  2a  of  these 
plates,  and  there  seems  to  be  no 
doubt  that  they  are  sclerotic  ele- 
ments. In  the  restoration  (fig.  26) 
29  sclerotic  plates  are  given,  but 
there  is  no  assurance  that  this 
number  is  the  exact  one.  They 
may  also  have  been  slightly  larger, 
but  not  as  large  as  in  Branchio- 
saurus. 

The  vertebral  column  is  pre- 
served nearly  entire  in  the  Daniels 
specimen  and  quite  entire  (478) 
in  the  Yale  specimen.  Cope,  in 
his  study  of  the  type  (105,  107), 
thought  there  could  be  no  more 
than  13  presacrals,  but  the  speci- 
men was  poorly  preserved  and  in- 
decisive on  this  point.  Dr.  Hay 
(316)  was  inclined  to  the  opinion 
that  there  were  less  than  20.  The 
Yale  specimen  shows  22  centra, 
which  are  elongate,  hour-glass- 
shaped  bodies,  with  the  neural 
spine  a  long,  low  crest  running 
the  entire  length  of  the  centrum, 


Fig.  26. — Restoration  of  body  outline  and  skeleton  of  .4  mphibamus 
grandiceps  Cope,  from  Mazon  Creek,  Illinois,  shales.  Restora- 
tion is  based  on  complete  specimens  of  the  species  and  on  Cope's 
drawing.  Form  of  body  is  indicated  in  one  specimen,  that  in 
possession  of  Mr.  Daniels.     X  1.5. 

Skull:  pmx,  premaxilla;  n,  nasal ;  fr,  frontal;  par,  parietal;  la,  lac- 
rimal; pf,  pref rental;  pof,  postfrontal;  po,  postorbital;  pp,  post- 
parietal;  spl,  supratemporal;  mx,  maxilla;j,  jugal;  qj,  quadrato- 
jugal;  sq,  squamosal;  tab,  tabulare. 

Skeleton:  ic,  interclavicle;  cl,  clavicle;  sc,  scapula;  h,  humerus;  r-u, 
radius,  ulna;  c,  carpus;  pu,  pubis;  il,  ilium;  /,  femur;  /,  tibia; 
fb,  fibula;  Is,  tarsus;  x,  ischium. 


with  a  median  elevation,  so  that  in  lateral  view  the  spine  would  be  triangular  in  form. 
The  body  of  the  centrum  is  expanded  laterally  into  a  diapophysis  which  extends 
anteriorly.     The  posterior  vertebrae,  at  least,  had  the  notochord  largely  persistent. 


THE   MICROSAURIAN   FAMILY  AMPHIliAMID^E. 


129 


t 


The  osseous  part  of  the  vertebra  seems  to  have  been  but  a  thin  shell,  and  the  struc- 
ture of  the  zygapophyses  can  not  be  determined.  That  they  were  dorsal  in  position 
is,  however,  evident  from  several  vertebrae.  The  points  of  these  structures  project 
laterally.     The  tail  is  short  and  the  caudal  vertebrae  weakly  developed. 

There  are  distinct  impressions  of  at  least  12  pairs  of  ribs  in  the  Daniels  specimen. 
They  are  long,  slender,  and  curved,  and  there  is  no  definite  assurance  that  there  were 
as  many  ribs  as  are  indicated  (fig.  26)  in  the  restoration  (462).  The  ribs  are  inter- 
central  (469)  and  probably  occu- 
pied the  full  length  of  the  verte-  *"^  ~r 
bral  column.  There  may  have 
been  as  many  as. indicated  in  the 
restoration. 

One  of  the  most  interesting 
features  of  the  Yale  specimen  is 
the  preservation  of  a  small  patch 
of  skin,  evidently  from  the  back, 
lying  to  one  side  near  the  head, 
measuring  5  mm.  in  length  by  3 
mm.  in  width.  The  fragment 
shows  the  skin  to  be  of  tubercu- 
lated  scales,  4  of  which  occupy 
the  length  of  1  mm.  The  scales 
are  somewhat  hexagonal,  almost 
rounded,  and  were  relatively 
quite  thick.  They  lie  in  a  close 
mosaic  (fig.  27). 

The  Yale  specimen  has,  very 
well  preserved,  a  portion  of  the 
ventral  scutelke,  of  the  throat, 
chest,  and  belly.  The  arrange- 
ment of  the  plates  on  the  throat 
and  chest  is  almost  exactly  the  reverse  of  what  Credner  has  described  (190)  for  Bran- 
ch iosa urus  amblystomus  Cred.  On  the  throat,  in  the  present  form,  the  chevron  points 
anteriorly,  and  it  is  the  anterior  prolongation  of  the  belly  scutes  with  the  postero- 
lateral projection  of  the  gular  scutes  which  form  the  chest  and  arm  scutellation.  The 
belly  chevrons  point  anteriorly,  as  in  Branchiosaurus,  the  rods  formed  by  the  scutes 
being  straight  and  not  curved  as  in  Bra nch iosa  urus .  The  entire  ventral  armature  pre- 
served is  displaced  to  the  left  of  the  animal  and  only  the  anterior  portion  is  preserved. 

The  pectoral  girdle  is  only  partially  known.  The  scapula  is  crescent-shaped. 
The  other  elements  are  indicated  only  by  fragments  and  nothing  is  known  of 
their  form. 

The  arm  elements  are  nearly  all  known.  The  humerus  is  slender  and  expanded 
at  the  ends,  with  its  articular  surfaces  well  developed.  The  separate  radius  and  ulna 
are  of  approximately  the  same  size  and  length.    The  carpus  is  unossified.    The  com- 


// 


^J 


A 


ts^- 


% 


Fig.  27. — Skeleton  of  Amphibamus  graiidiceps  Cope.     X  i-4- 

carpus;  cl,  clavicle;  cr,  caudal  rib;  cv,  caudal  vertebra;/,  femur; 
h,  humerus;  il,  ilium;  s,  skin;  or,  orbit;  r,  radius;  «,  ulna;  sc, 
scapula;  sp,  sclerotic  plates;  /,  tibia  and  fibula;  Is,  tarsus;  vs, 
ventral  scutelte.     Specimen  No.  794,  Yale  University  Museum. 


I30  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

plete  phalangeal  formula  for  the  hand  of  Amphibamus  is  unknown.  The  third 
digit  seems  to  have  4  elements.    The  formula  2-2-3-2  has  been  suggested  (462). 

The  pelvis  is  very  satisfactorily  known.  The  ilium  is  a  long,  slender,  straight 
rod,  with  expanded  ends.  The  ischium  is  shown  on  both  sides  of  the  vertebral 
column  in  the  Yale  specimen.  Its  form  is  almost  identical  with  that  of  Paleohatteria 
longicaudata  Credner,  from  the  Rothliegenden  of  Saxony.  The  ischia  are  apparently 
approximate  in  the  median  line,  though  this  character  is  somewhat  obscured  by  the 
impression  of  the  caudal  vertebrae.  Their  relation  with  the  ilium,  other  than  that 
they  were  posterior  to  it,  is  uncertain.  The  pubis  is,  apparently,  calcified  cartilage. 
It  is  a  squarish  plate,  somewhat  corrugated,  lying  anterior  to  the  ilium  in  the  Dan- 
iels specimen.  The  elements  of  the  pelvis  were  undoubtedly  hung  loosely  in  the 
flesh,  as  in  modern  salamanders,  since  there  is  no  indication  of  articular  surfaces. 

The  hind  limb  is  well  known,  the  type  having  a  nearly  complete  leg  with  the 
foot.  The  Daniels  and  the  Yale  specimens  supplement  and  substantiate  the  type. 
The  femur  is  longer  than  the  humerus,  but  more  slender,  with  its  articular  surfaces 


Fig.  28. — Restoration  of  probable  appearance  of  Amphibamus  grandiceps  Cope  on  the 
basis  of  the  material  described  herewith.     X  1.5. 

about  as  well  developed  as  in  the  humerus.  The  element  is  a  simple  rod  of  bone 
without  muscular  crests  of  any  kind.  The  tibia  and  fibula  are,  likewise,  slender 
separate  rods  of  bone.  The  tarsus  is  unossified.  The  phalangeal  formula  is  2-2-3- 
4-3,  and  is  fairly  definite. 

In  the  type  specimen  the  matrix  in  the  orbit  was  blackened  as  if  by  the  pigmen- 
tum  nigrum  of  the  choroid.  The  same  has  been  noticed  in  other  specimens.  Pro- 
fessor Cope  thought  this  indicated  that  the  animal  was  nocturnal. 

There  are  many  characters  in  Amphibamus  which  seem  to  approximate  the  rep- 
tilian type  of  structure.  Among  these  may  be  mentioned  the  character  of  the  artic- 
ular surfaces  of  the  limb  bones,  the  intercentral  position  of  the  ribs,  the  incipient 
double-headedness  and  the  curvature  of  the  ribs,  the  presence  of  a  cartilaginous 
calcified  pubis,  the  length  of  the  limbs,  and  the  clawed  character  of  the  phalanges. 

Amphibamus  was  a  low,  flat,  short,  and  undoubtedly  a  creeping,  crawling  ani- 
mal, possibly  spending  a  portion  of  its  time  in  the  water;  but  it  could  not  have  been 
a  swimmer.    It  was  one  of  nature's  first  attempts  at  constructing  a  land  vertebrate. 


THE   MICROSAURIAN   FAMILY   AMPHIBAMID.E. 


131 


Measurements  of  Amphibamus  ukandiceps  Cope. 


Collection  of  Mr.  L.  E.  Daniels,  of  Rolling 
Prairie,  Indiana: 

mm. 

Entire  length  of  specimen 62 

Posterior  width  of  head 15 

Length  of  head 15 

Posterior  height  of  skull 3 

Length  of  orbit 5 

Width  of  orbit 3.5 

Interorbital  width 4 

Width  of  skull  in  front  of  orbits n 

Width  of  skull  just  back  of  orbits 16 

Length  of  presacral  region  of  the  vertebral 

column 30 

Length  of  tail 13 

Length  of  fore  limb 13.5 

Length  of  humerus 4 

Length  of  radius  and  ulna 3 

Length  of  right  hand  as  preserved 3.5 

Length  of  rib  along  curve 5.5 

Length  of  hind  limb 17 

Length  of  ilium 4 

Length  of  vertebral  centrum 1 .75 

Length  of  portion  of  scapula  (?)  preserved.  4.5 

Length  of  foot 6.5 

Width  of  impression  of  body  midway 16 


No.  794  (1234),  Yale  University  Museum: 

mm. 

Length  of  skeleton 67 

Length  of  skull 15 

Posterior  width  of  skull 15 

Depth  of  tympanic  notch 4 

Width  of  tympanic  notch 6 

Long  diameter  of  the  orbit 7 

Transverse  diameter  of  the  orbit 5.5 

Interorbital  width 4.5 

Diameter  of  pineal  foramen .75 

Length  of  cervical  series  of  vertebra? 9 

Length  of  dorsal  series 35 

Length  of  caudal  series 13 

Length  of  a  centrum  of  the  dorsal  series ....        1 .5 

Length  of  dorsal  rib 3.5 

Length  of  arm 20 

Length  of  humerus 7 

Length  of  radius  and  ulna 4 

Width  of  carpal  space 3 

Length  of  third  digit 5 

Length  of  leg 25 

Length  of  ilium 3 

Length  of  femur 9 

Length  of  tibia  and  fibula 5 

Length  of  carpal  space 4 

Length  of  1st  digit 3 

Length  of  2d  digit 4.5 

Length  of  4th  digit 7 

3  ventral  scutellae  in  1  mm. 

Amphibamus  thoracatus  Moodie. 

Moodie,  Proc.  U.  S.  Nat.  Mus.,  40,  pp.  431-433,  fig.  2,  191 1. 

Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  pp.  347-349,  pi.  5,  fig.  2,  1912. 

Type:  Specimen  No.  4306,  U.  S.  National  Museum. 

Horizon  and  locality:  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  type  is  a  part  of  the  collection  of  Mr.  R.  D.  Lacoe,  in  the  U.  S.  National 
Museum.  The  fossil  is  very  poorly  preserved,  but  the  remains  are  to  be  seen  on  both 
halves  of  the  nodule,  so  that  considerable  can  be  made  out  as  to  its  structure. 

The  chief  diagnostic  characters  which  will  at  once  distinguish  the  species  are  the 
elongate  arm,  large  interclavicle,  shape  of  the  vertebra,  and  triangular  skull. 

The  portions  of  the  animal  which  are  preserved  are  the  impression  of  the  skull 
with  one  orbit,  the  right  humerus  and  radius  with  portions  of  others,  and  traces  of 
ventral  scutellae.  These  remains  are  so  intermingled  with  the  remains  of  plants 
that  it  has  been  quite  difficult  to  distinguish  bone  impression  from  plants.  This, 
however,  has  been  done  by  whitening  the  fossils  with  ammonium  chloride,  when  the 
texture  of  the  fossils  serves  to  distinguish  the  one  from  the  other.  Parts  of  the  plants 
have  been  converted  into  galena  and  kaolin,  as  have  also  parts  of  the  bones,  so  the 
task  has  been  rendered  doubly  difficult.  There  can  be  no  doubt,  however,  that  the 
observations  recorded  below  are  correct.  The  position  of  the  arm  in  relation  to  the 
pectoral  girdle  and  the  position  of  the  girdle  in  relation  to  the  skull  impression  first 
called  attention  to  the  possible  presence  of  a  fossil  amphibian. 

There  is  little  to  be  said  of  the  skull.  It  is  merely  an  impression  in  the  nodule. 
It  is  triangular  in  form,  with  the  snout  an  acute  angle.  The  angle  is,  however,  exag- 
gerated by  the  compression  to  which  the  fossil  has  been  subjected.  The  right  side  of 
the  skull  lies  over  a  portion  of  some  plant.  The  animal  is  preserved  on  its  back,  so 
that  this  gives  a  good  opportunity  for  the  study  of  the  pectoral  girdle,  which  is  par- 


132  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

tially  preserved.  The  interclavicle  is  very  large  and  from  it  the  species  has  been 
given  its  specific  name  {thoracatus — armed  with  a  breast  plate) .  It  is  an  exaggerated 
T,  with  the  stem  very  short  with  its  anterior  margin  curved,  and  ending  in  a  rather 
sharp,  elongate  point.  The  interclavicle  recalls,  in  a  measure,  the  same  element  of 
the  Branchiosauria,  although  it  is  much  more  expanded  anteriorly  and  has  a  shorter 
spine.    In  these  respects  it  resembles  more  nearly  a  reptilian  interclavicle  (fig.  14  B) . 

The  clavicle  is  of  the  simple  triangular  shape  so  characteristic  of  the  Microsauria. 
It  is  somewhat  displaced  backward  and  its  inner  margin  is  slightly  obscured.  The 
humerus  is  elongate,  apparently  cylindrical,  and  with  expanded  ends,  resembling 
very  closely  the  humerus  of  Amphibamus  grandiceps,  although  its  proportions  are 
much  greater  than  in  that  species.  Its  length  is  almost  equal  to  the  length  of  the 
skull,  while  in  A.  grandiceps  the  length  of  the  humerus  is  only  half  that  of  the  skull. 
The  radius  (ulna?)  resembles  in  its  general  proportions  those  of  the  humerus.  It  is  a 
more  elongate,  slender,  lighter  bone.  The  impression  of  the  other  bone  of  the  fore- 
arm is  obscured. 

A  portion  of  a  single  vertebral  centrum  from  the  posterior  part  of  the  dorsal 
series  is  preserved.  It  is  apparently  amphiccelous ;  its  width  is  nearly  half  greater 
than  its  length. 

Measurements  of  the  Type  of  Amphibamus  thoracatus  Moodie. 
(No.  4306,  U.  S.  National  Museum.) 

mm.  mm. 

Length  of  entire  specimen,  as  preserved 60  Greatest  transverse  diameter 3 

Length  of  skull  impression 18  Length  of  humerus 10 

Greatest  width  of  same 15.5  Greatest  diameter  ( >f  same 4 

Long  diameter  of  right  orbit .  4  Least  diameter  of  same 1.5 

Transverse  diameter  of  same 3  Length  of  radius  (ulna?) 11 

Transverse  width  of  interclavicle 14  Length  of  vertebral  centrum 2 

Long  diameter  of  same 7(?)  Width  of  same 3 

Long  diameter  of  clavicle 9 

Genus  CEPHALERPETON  Moodie. 

Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  349,  1912. 

Type:  Cephalerpeton  ventriarmatum  Moodie. 

This  genus  is  founded  on  remains  of  a  nearly  entire  individual  of  a  relatively 
large  microsaurian  from  the  Mazon  Creek  shales.  The  genus  is  most  immediately 
related  to  the  Amphibamida?,  of  which  two  species  are  already  known,  Amphibamus 
grandiceps  Cope  and  A.  thoracatus  Moodie.  The  present  genus  differs  from  these 
species  in  many  respects,  notably  in  size.  The  skull  in  Cephalerpeton  is  nearly  as 
long  as  half  the  entire  body  of  Amphibamus  grandiceps  Cope,  inclusive  of  the  tail. 
Other  structural  differences  are  the  anisodont  teeth,  the  large  size  and  the  more 
median  position  of  the  orbits,  and  the  absence  of  the  posterior  tympanic  notch 
in  Cephalerpeton.  The  form  of  the  skull  recalls  that  of  Melanerpeton  and  Pclo- 
saurus  (190)  of  Europe,  but  those  genera  are  branchiosaurian,  while  the  present 
form,  from  the  structure  of  the  vertebrae  and  the  long,  curved  ribs,  is  an  un- 
doubted microsaurian.  Nothing  like  it  occurs  in  any  of  the  amphibian  faunas 
thus  far  made  known.  It  is  most  nearly  approached  by  a  member  of  the  genus 
Erpetosaurus,  but  from  this  genus  the  present  form  is  readily  distinguished  by  the 
smooth  skull  bones,  the  absence  of  a  posterior  table  to  the  skull,  and  the  presence 
of  a  highly  developed  ventral  armature.  The  interorbital  width  is  less  than  the 
transverse  diameter  of  the  orbit. 


PLATE   18 


&A 


4. 


Type  specimen  of  Erpetosanrtis  sailptilis  Moodie,  from  the  Cannelton  Shales  of  Pennsyl- 
vania.    Original  in  the  University  of  Chicago,  Walker  Museum. 

Skeletal  elements  of  Eryops  sp.  indet.,  from  the  Pittsburgh  Red  Shale  at  Pitcairn, 
Pennsylvania.  <7=nearly  complete  vertebra;  b  and  r=ribs;  </=pleurocentrum; 
f=neural  arch  and  spine.  Originals  in  the  Carnegie  Museum  at  Pittsburgh. 
After  Case. 

Photograph  of  amphibian  footprints,  Dromoput  adiimus  Branson,  from  the  Mississippian 
shales  of  Giles  County,  Virginia.  X  %.  Courtesy  of  Dr.  Branson.  Original  in  the 
Museum  at  Oberlin  College. 

Photograph  of  type  of  V'/iim/piis  antitjiius  Marsh,  the  amphibian  footprint  from  the 
Devonian  of  Pennsylvania.  X  %.  Courtesy  of  Dr.  Lull.  Original  No.  784,  Vale 
University  Museum. 


THE   MICROSAURIAN   FAMILY  AMPHinAMID.4i. 


133 


Cephalerpeton  ventriarmatum  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  pp,  350-352,  pi.  I,  fig.  4;  pi.  7,  fig.  2,  1912. 

Type:  Specimen  No.  796,  of  Yale  University  Museum. 

Horizon  and  locality:  Collected  at  Mazon  Creek  in  1871,  near  Morris,  Illinois 

The  remains  on  which  the  present  species 
is  based  consist  of  an  almost  entire  skull,  26 
consecutive  vertebrae,  both  fore  limbs,  20  ribs 
preserved  on  the  right  side  of  the  body,  and  a 
portion  of  the  ventral  armature  (plate  4,  fig.  4). 

The  skull  is  very  broad  posteriorly,  its 
width  being  one-third  greater  than  its  length, 
with  due  allowance  for  crushing.  A  pineal 
foramen  is  not  preserved.  The  sutures  bound- 
ing the  premaxillaries,  the  maxillae,  the  nasals, 
the  prefrontals,  the  frontals,  a  portion  of  the 
parietals,  the  squamosal,  the  supratemporal, 
the  quadratojugal,  and  the  quadrate  (?)  are 
fairly  well  preserved.  The  arrangement  of 
these  elements  can  be  discerned  by  reference 
to  figure  29.  The  prefrontals  are  unusually 
large  and  are  triangular  in  shape.  The  supra- 
temporal  is  also  quite  large.  The  surface  of 
the  skull  bones  is  smooth  and  there  is  nowhere 
an  indication  of  sculpture. 

Portions  of  4  sclerotic  plates  are  preserved 
in  the  right  orbit.  These  measure  0.5  by  0.75 
mm.  The  orbits  are  large  and  the  interorbital 
space  is  less  than  the  transverse  diameter  of 
the  orbit.  Thirteen  teeth,  apparently  pleuro- 
dont,  are  preserved  on  the  left  maxilla.  They 
are  short,  sharply  pointed,  smooth,  and  unequal.  The  first  2  left  maxillary  teeth 
from  the  anterior  end  are  short;  then  follows  a  tooth  which  is  one-third  longer 
than  these  two;  the  fourth  tooth  is  somewhat  shorter  than  the  third;  the  fifth 
and  sixth  are  still  shorter  and  are  practically  equal  in  size,  though  somewhat  larger 
than  the  first  two. 

The  right  mandible  is  preserved  almost  entire,  though  so  badly  eroded  that  little 
can  be  said  of  its  structure.  Impressions  of  1 2  teeth  are  present  on  the  mandible  and 
all  are,  apparently,  equal.  The  cotylus  seems  to  have  been  far  posterior  and  an 
angle  of  the  mandible  projected  slightly  back  of  the  skull. 

There  remain  only  a  few  indefinite  impressions  of  the  cervical  vertebrae.  The 
union  of  the  skull  with  the  vertebral  column  is  obscured  and  lost.  Impressions  of 
the  dorsal  vertebrae  are  well  preserved,  and  wax  molds  made  from  these  show 
the  structure  of  the  dorsal  vertebrae  surprisingly  well.  They  are  long  and  cylin- 
drical, with  the  median  portions  slightly  constricted  by  a  deep  pit  on  each  side  of 


Fig.  29. — Skeleton  of  Cephalerpeton  ventriarmatum 
Moodie.     X  I. 

pf,  prefrontal ;  d,  clavicle ;  m,  mandible ;  h,  humerus ; 
j,  jugal;  mx,  maxilla;  or,  orbit;  ph,  phalanges 
of  hand;  par,  parietal;  po,  postorbital;  r, 
radius;  sp,  sclerotic  plates;  u,  ulna;  vs,  ventral 
scutella;. 


134  THE    COAL    MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

the  low  neural  ridge,  which  takes  the  form  observed  in  Thyrsidium,  Molgophis,  Phleg- 
ethontia,  Dolichosoma  (fig.  8)  and  other  genera.  The  vertebrae  are  strongly  amphi- 
ccelous  and  the  notochord  was  probably  persistent.  The  sides  of  the  vertebras  are 
smooth. 

The  ribs  are  all  intercentral  in  position;  the  anterior  ones  very  broad  near  the 
base,  recalling  the  broadly  expanded  ribs  described  by  Schwarz  (540)  for  Scincosau- 
rus,  Ptyonius,  Thyrsidium,  and  other  genera.  Posteriorly  the  ribs  become  slender 
and  cylindrical.  They  are  all  rather  long  and  distinctly  curved,  with  probably  a 
cartilaginous  tip. 

There  is  preserved  a  single  element  of  the  right  side  of  the  pectoral  girdle.  This 
is,  I  think,  the  coracoid,  an  element  which  has  hitherto  escaped  observation  among 
the  American  Microsauria.  It  is  long  and  spatulate  at  both  ends,  with  the  median 
portion  apparently  almost  cylindrical,  not  unlike  that  described  by  Credner  (181) 
for  the  coracoid  of  Branchiosaurus,  save  that  the  lower  end  of  the  branchiosaurian 
coracoid  is  acuminate.  In  the  present  form  it  is  spatulate.  Its  relations  with  the 
other  elements  of  the  pectoral  girdle  have  never  been  satisfactorily  determined. 

The  fore  limbs  are  both  partially  preserved.  The  humerus  of  the  right  side  is 
complete.  It  is  greatly  elongated  for  a  microsaurian.  The  form  of  the  element  is 
not  unlike  that  of  a  lizard,  with  the  lower  end  of  the  bone  spatulate  and  endochon- 
drium  well  developed.  Very  little  difference  can  be  seen  between  the  form  of  the 
arm  bones,  which  represent  the  radius  and  ulna.  They  are  both  elongated,  with  con- 
stricted median  portion  and  expanded  truncate  ends.  The  carpus  is  unossified  and 
the  cartilage  has  left  no  trace  of  the  elements. 

The  right  hand  has  two  metacarpals  preserved,  which  are  fully  half  as  long  as 
the  radius  and  ulna.  They  are  separated  some  little  distance  from  the  ends  of  these 
elements,  though  this  may  be  due  to  post-mortem  shifting.  The  carpus  may,  how- 
ever, have  been  broad.  On  the  left  side  are  preserved  portions  of  the  humerus, 
radius,  ulna,  and  3  metacarpals,  lying  close  to  the  vertebral  column.  The  carpal 
space  is  not  so  large  on  the  left  as  on  the  right.  The  ventral  armature  is  well 
preserved  in  a  narrow  patch  about  an  inch  in  length.  The  chevron-shaped  rods 
are  quite  large,  there  being  2  of  them  in  1  mm. 

Measurements. 

mm.  mm. 

Entire  length  of  fossil 98  Median  width  of  a  centrum 1.5 

Length  of  skull 22  Length  of  rib 6.5 

Width  across  base  of  skull 28  Width  of  rib  at  base 33 

Long  diameter  of  eye 10.5  Length  of  coracoid 8 

Transverse  diameter  of  eye 8  Width  of  coracoid  at  anterior  end 2.5 

Interorbital  space 4  Length  of  carpal  space 5 

Length  of  mandible 26  Length  of  humerus 18 

Depth  of  mandible  at  coronoidal  region 3.5  Width  of  shaft I 

Depth  of  dentary 2  Distal  width  of  humerus 4 

Length  of  long  tooth 2  Length  of  radius  and  ulna 10.5 

Diameter  of  long  tooth  at  base .5  Length  of  metacarpal 6 

Length  of  preserved  portion  of  vertebral  column ...  64  Length  of  ventral  armature  preserved 24 

Length  of  a  centrum 3  Number  of  rods  in  length  of  5  mm 10 


Type  specimen  of  Ctenerpeton  alveolatum  Cope,  from  the  Coal  Measures  of  Ohio.     X  1.33. 
Original  in  U.  S.  National  Museum. 


MOOOIf 


Skull  of  Erpetosaurus  minutus  Moorlie,    from   the   Cannelton   slates   of     Pennsylvania. 

Original  in  U.  S.  National  Museum.     Enlarged  X  3.3. 
Skull  and  anterior  part  of  body  of  Ptyonhis  ftcclinatus  Cope,  from  the  Coal    Measures   of 

Linton,  Ohio.     Original  in  U.  S.  National  Museum.     X  1. 
Skeleton  of' Eosauravus  copei  Williston,  from  the  Coal  Measures  of  Linton,  Ohio.     "The 

oldest  known  reptile  from  North  America"  and  closely    related   structurally    to   the 

Microsauria.     Original  in  U.  S.  National  Museum.     X  1. 
l'art  of  the  ventral  scutellation  and  ribs  of   Sauroplevra  iti»i/ata    Cnpe,    from    the    dial 

Measures  of  Linton,  Ohio.   Original  in  American  Museum  of  Natural  History.      X  1. 


CHAPTER  XVIII. 

THE  M1CROSAURIAN  FAMILY  NYRAN1ID/E,  FROM  THE  COAL  MEASURES  OF  OHIO. 

Family  NYRANIID.E  Lydekker,  1890. 

Lydekker,  Cat.  Fossil  Reptilia  and  Amphibia,  p.  166,  1890. 

Skull  with  the  palatines  situated  near  the  middle  line,  internally  to  the 
vomers  and  pterygoids,  and  the  palatine  vacuities  small  and  placed  far  back. 
Vertebras  (Ichthyerpeton)  discoidal.  Teeth  less  complex  than  in  the  Anthraco- 
sauridae.  A  ventral  armor  present  and  the  entire  body  covered  with  small  cycloid 
imbriated  scales. 

The  type  genus  of  this  family  was  placed  by  Fritsch  (251)  with  the  Archegosau- 
ridae,  although  its  resemblance  to  Anthracosaurus  was  pointed  out;  it  was  subse- 
quently made  the  type  of  a  family  by  Lydekker  (393)  in  1890,  and  placed  next  the 
Archegosauridae.    Known  from  the  Coal  Measures  of  Bohemia,  Ireland,  and  Ohio. 

Two  genera  from  North  America,  Ichthyerpeton  and  Cer car iomor phis,  are 
assigned  tentatively  to  this  family,  both  known  from  the  Coal  Measures  (462)  of 
Linton,  Ohio,  and  both  with  the  body  completely  scaled.  The  distinguishing  char- 
acters are  found  chiefly  in  the  shape  and  arrangement  of  the  scales,  the  structure, 
form,  and  size  of  the  body,  all  of  which  are  given  full  treatment  in  the  discussion 
below. 

Genus  ICHTHYERPETON  Huxley,  1866. 

Huxley,  Trans.  Roy.  Irish  Acad.,  xxiv,  p.  195,  pi.  xxiii,  fig.  1;  Scientific  Memoirs,  in,  p.  195,  pi.  23,  fig.  1, 
1866. 

The  genus  was  founded  by  Huxley  (334)  for  the  reception  of  the  species  Ichthy- 
erpeton bradleyce  from  the  Kilkenny  Coal  Measures  of  Ireland.  The  remains  of  the 
type  specimen  represent  "the  hinder  moiety  of  the  trunk,  with  the  greater  part  of 
the  tail,  of  an  animal  whose  scaly  integument  and  laterally  compressed,  fin-like  tail 
might  easily  lead  one  to  take  it  for  a  fish,  were  not  its  true  position  among  higher  ver- 
tebrata  settled  at  once  by  the  digitate  hind  limb ;  while  its  alliance  with  the  labyrin- 
thodonts  is  indicated  by  the  delicate  spicular  ossicles,  which  form  a  rudimentary 
dermal  shield  along  the  belly."  (Huxley.) 

Ichthyerpeton  squamosum  Moodie. 

Moodie,  Jour.  Geol.,  XVII,  No.  I,  p.  69,  1909. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  24,  1909. 

Type:  Specimens  Nos.  4476  and  4459,  U.  S.  National  Museum. 

Locality  and  horizon :  Linton,  Ohio,  Coal  Measures. 

The  present  species  is  based  on  well-preserved  remains  from  the  Linton,  Ohio, 
beds.  There  are  two  specimens  of  the  species  preserved  on  blocks  of  coal  and 
together  they  represent  the  greater  part  of  the  length  of  the  animal.  The  species  is 
located  in  the  genus  Ichthyerpeton,  which  was  founded  by  Huxley  (334,  p.  351)  on 
remains  from  the  Coal  Measures  of  Ireland,  on  account  of  the  character  of  the  der- 

135 


!36  THE  COAL  MEASURES  AMPHIBIA  OF  NORTH  AMERICA. 

mal  covering,  which  consists  of  small  scales  such  as  Huxley  described  in  the  form 
from  Ireland.  The  specific  characters  of  this  form  are  the  small  size  of  the  rounded 
scales,  the  attenuated  tail,  the  apparent  absence  of  limbs,  the  character  of  the  ven- 
tral scutellation,  and  the  slightly  curved  condition  of  the  ribs. 

It  is  estimated,  from  the  portions  preserved,  that  the  animal  attained  a  length 
of  not  less  than  3  feet  and  its  body  was  long  and  slender.  It  may  have  had  an 
appearance  similar  to  the  modern  caudate  genus  Siren,  though  there  were  doubtless 
4  limbs  present  instead  of  2.  The  slenderness  of  the  body  is  at  variance  with  the  con- 
dition found  in  the  type  species  Ichthyerpeton  bradleyce  Huxley,  in  which  the  trunk 
was  rather  stoutly  built.  The  character  of  the  anterior  portion  of  the  body  in  the 
present  species  can  not  be  determined  and  the  skull  is  wanting.  There  are  no  evi- 
dences of  anterior  limbs,  although  the  ventral  scutellation  preserved  would  seem  to 
include  the  pectoral  region.  No  pectoral  shields  are  preserved,  nor  are  there  any 
traces  of  pelvic  girdle  or  limbs. 

The  preserved  portions  on  one  block  include  nearly  the  entire  tail  and  the  pos- 
terior region  of  the  body,  and  on  the  other  block  the  dorsal  region  of  the  body  and 
the  anterior  portion  of  the  tail,  so  that  the  two  specimens  supplement  each  other 
in  an  interesting  manner.  There  are  impressions  of  several  vertebras  preserved. 
They  are  much  the  same  in  character  as  Huxley  has  described  for  the  type  species 
(I.  bradleyce).  They  are  short  and  thick  and  were  probably  amphiccelous.  There 
are  likewise  preserved  the  remains  of  rather  slender  recurved  ribs  mingled  in  with 
the  remains  of  the  ventral  scutellation  and  distinguished  from  the  elements  of  the 
abdominal  shield  by  their  size  and  curvature.  They  are,  apparently,  single-headed, 
but  the  character  of  their  articulation  can  not  be  determined.  The  ventral  scutel- 
lation consists  of  fine  continuous  rods  arranged  in  the  regular  chevron  pattern. 
They  do  not  seem  to  be  divided  into  oat-shaped  scutes,  as  is  the  case  with  the  form 
described  by  Huxley.  The  ventral  rods  are  closely  packed  for  a  distance  of  more 
than  6  inches,  but  as  they  are  scattered  their  exact  arrangement  can  not  be  deter- 
mined. They  seem  to  have  extended  to  the  cloacal  region,  but  there  are  no  evidences 
of  the  specialized  clasping  organs  such  as  Fritsch  (251)  has  described  in  the  ven- 
tral scutellae  of  Ophiderpeton.  The  scales,  which  are  well  preserved  on  the  tail,  may 
have  covered  the  entire  body,  since  there  are  many  scattered  scales  in  the  dorsal 
region  of  one  of  the  specimens.  They  are  slightly  oval,  tuberculate,  and  measure 
scarcely  1  mm.  in  their  longest  diameter.  They  show  but  slight  evidences  of  having 
been  imbricated,  though  it  is  likewise  possible  that  they  were  simply  inclosed  within 
the  integument,  and  somewhat  separated  from  one  another.  The  most  posterior 
part  of  the  tail  preserved  seems  to  indicate  that  the  tip  was  attenuated.  It  was  prob- 
ably flattened  from  side  to  side.  We  may  thus  regard  Ichthyerpeton  squamosum  as 
an  elongate  aquatic  animal  with  a  long,  flattened  tail,  and  since  there  were  possibly 
no  limbs  or  very  small  ones,  it  would  be  an  animal  highly  adapted  for  life  in  the 
water.  The  present  species  is  of  interest  because  it  represents  an  additional  dis- 
covery of  the  scaled  Amphibia  in  North  America.  The  species  previously  known 
from  the  Linton,  Ohio,  deposits  is  Cercariomor phus  parvisquamis  Cope.  Dermal 
scales  have  also  been  observed  in  specimens  of  Amphibamus  grandiceps  Cope  and 


1. 


4. 

5. 
6. 


Mandible  of  Macrerpeton  deani  Moodie,  from  the  Linton,  Ohio,  Coal  Measures. 
Original  in  American  Museum  of  Natural  History,  No.  2934.     X  0.6. 

Portion  of  the  skull  of  Macrerpeton  deani  Moodie,  possibly  of  the  same  individual 
as  the  mandible.  From  the  Linton,  Ohio,  Coal  Measures.  Original  in 
American  Museum  of  Natural  History,  No.  8535  G.     X  0.4. 

Type    of    CetvariemorphtU    pandsquamis    Cope,     from    the  Linton,    Ohio,    Coal 

Measures.     Original  in  American  Museum  of  Natural  History.     X  1. 
An  additional  specimen  of  Cercariomoiphus  pa/i'isi/naiiiis  Cope,  from  the    Linton, 

( >hio,  Coal  Measures.  Originalin  American  Museum  of  Natural  History.  X  1. 
Skull  of  Sauroptcnra  scntcllata  Newberry.  From  the  Coal  Measures  of  Ohio.  X  1. 
Tooth  of  Mastodonsaurus  sp.  indet.  of  the  Carboniferous  of    Kansas.     Original   in 

University  of  Kansas  Museum.     X  1. 
Tooth    of    Mastodonsaurus    °ivan/eus    Jaeger,    from    the    Triassic    of    Germany. 

Introduced  for  comparison  with  the  tooth  from  the  Kansas  Carboniferous.     XI. 


THE    MICROSAURIAN   FAMILY   NYRANIIDjE.  1 37 

Micrerpeton  caudatum  Moodie  (462,  478)  from  the  Mazon  Creek,  Illinois,  beds,  and 
Sir  William  Dawson  (208)  described  scales  accompanying  several  forms  from  the 
Joggins  deposits  of  western  Nova  Scotia. 

Measurements  of  the  Types  of  Ichthyerpeton  squamosum  Moodie. 

Length  of  animal  as  estimated  from  two  impressions 3  ft. 

Length  of  longest  impression 21  in. 

Length  of  specimen  containing  tail  impression 9  in. 

Width  of  tail  impression:     Maximum 50  mm. 

Minimum 6  mm. 

Width  of  a  single  scale 1  mm. 

Distance  from  base  of  tail  to  tip 125  mm. 

Length  of  specimen  as  preserved 225  mm. 

Width  of  chevron  rod  space 30  mm. 

Length  of  rib 25  mm. 

8  chevrons  in  a  distance  of  3  mm. 

Genus  CERCARIOMORPHUS  Cope. 

Cope,  Proc.  Amer.  Phil.  Soc,  1885,  p.  405. 

Type:  Cercariomorphus  parvisquamis  Cope. 

The  type  specimen  of  this  genus  is  supplemented  by  a  portion  of  the  body  of 
another  specimen  which  adds  a  little  to  our  knowledge  of  the  animal's  form,  but 
nothing  as  to  structure.    Cope's  original  description  is  as  follows : 

"Represented  by  a  fusiform  body  which  terminates  in  a  long,  slender,  cylindrical  tail, 
and  which  is  covered  with  small  subquadrate  scales  quincuncially  arranged.  No  fins  or 
limbs  are  preserved,  and  the  form  of  the  head  can  not  be  made  out.  Probably  a  portion 
of  the  skull  is  preserved.  There  are  some  scattered  bodies  in  the  body  portion,  which 
look  like  deeply  concave  vertebrae  with  the  zygapophyses  of  batrachians.  There  are  some 
linear  impressions  at  one  point,  which  resemble  the  bristle-like  rods  on  many  Stegocephali. 
They  are  so  few  as  to  be  of  little  importance.  The  scales  are  like  those  of  fishes.  There 
are  traces  of  segmentation  in  the  axis  of  the  long  tail. 

"The  position  of  this  curious  form  is  quite  uncertain.  It  is  quite  different  from  any- 
thing observed  hitherto  in  the  American  Coal  Measures." 

Cercariomorphus  parvisquamis  Cope. 

Cope,  Proc.  Amer.  Phil.  Soc,  1885,  p.  405. 
Moodie,  Science,  n.s.,  xli,  No.  1056,  p.  463,  1915. 

Type :  Specimen  No.  2560,  Newberry  Collection,  American  Museum  of  Natural 
History. 

Horizon  and  locality:  Discovered  by  Samuel  Huston  at  the  Linton,  Ohio,  Coal 
Mines.     (Plate  21,  figs.  3,  4;  24,  figs.  2,  3.) 

The  scales  (plate  24,  fig.  2)  in  their  present  condition  are  entirely  smooth.  At  a 
distance  of  20  mm.  from  the  base  of  the  tail  they  are  in  20  longitudinal  series.  At 
that  point  the  transverse  diameter  of  the  body  is  140  mm.  The  outline  contracts 
rather  abruptly  to  the  tail,  of  which  66  mm.  are  preserved.  The  surface  of  the  tail  is 
obscured  by  a  thin  layer  of  carbonaceous  matter  not  sufficiently  thick  to  obscure 
scales,  which  are  evident  at  distances  of  16  mm.,  43  mm.,  and  52  mm.  from  the  tip. 
The  scales  on  the  tail  are  smaller  than  those  on  the  body  and  are  without  markings 
of  any  kind.  The  anterior  half  of  the  body  is  depressed  and  distorted,  but  the 
remainder  is  well  preserved  and  shows  a  fairly  good  outline  of  an  apparently 
limbless  body. 


I38  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

An  additional  specimen  (No.  8683  G,  of  the  Newberry  Collection,  American 
Museum  of  Natural  History)  reveals  no  new  facts  as  to  structure,  but  serves  to 
show  that  the  body  of  the  animal  was  long  and  slender  (plate  21,  fig.  4).  The  por- 
tion studied  comes  undoubtedly  from  the  middle  of  the  body.  No  limb  elements 
are  preserved.  The  scales  are  somewhat  larger,  especially  toward  the  sides  of  the 
body,  than  in  the  type.  The  fragment  measures  70  mm.  in  length  by  18  mm.  and 
26  mm.  in  width.    One  of  the  largest  scales  measures  1  mm.  in  diameter. 

Measurements  of  Type. 

mm. 

Length  of  entire  remains 180 

Greatest  width 22 

Greatest  width  of  undisturbed  portion 15 

Length  of  an  individual  scale .75 


CHAPTER  XIX. 

THE  AISTOPODOUS  MICROSAURIAN  FAMILY  PTYONIID/E,  FROM  THE  COAL 

MEASURES  OF  OHIO. 

Family  PTYONIID^  Cope,  1875. 

Cope,  Gcol.  Surv.  Ohio,  11,  pt.  11,  p.  357,  1875. 

Elongate,  slender,  weak-limbed,  aquatic  microsaurians.  Neural  and  haemal 
spines  of  vertebra  elongated,  expanded  and  sculptured.  Ventral  armature  weakly 
developed  or  absent.    Skull  lanceolate,  with  long,  slender  teeth. 

Three  genera  are  assigned  to  this  family:  Ptyonius,  (Estocephalus,  and  Thyr- 
sidium.  The  forms  are  very  closely  related,  and  when  additional  material  is  secured 
the  three  genera  may  be  found  to  be  identical.  The  species  included  in  this  family 
are:  Ptyonius  pectinatus  Cope,  P.  vinchellianus  Cope,  P.  marshii  Cope,  P.  nummifer 
Cope,  P.  serrula  Cope,  (Estocephalus  remex  Cope,  0.  rectidens  Cope,  Thyrsidium  fas- 
ciculare  Cope.  The  species  are  all  exclusively  from  the  Linton,  Ohio,  Coal  Measures, 
and  most  of  them  are  known  from  abundant  material. 

Genus  PTYONIUS  Cope,  1875. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  373,  1875. 

Cope  designated  no  species  as  the  type,  but  we  may  regard  Ptyonius  pectinatus 
as  typical. 

Form  elongate,  with  long  tail  and  lanceolate  cranium.  Limbs  weak,  a  posterior 
pair  only  discovered.  Three  clavicular  elements;  abdomen  protected  by  packed 
osseous  rods,  which  are  arranged  en  chevron,  the  angle  directed  forward.  Neural 
and  haemal  spines  of  caudal  vertebrae  expanded  and  fan-like.  Ribs  well  developed. 
The  various  species  vary  in  length  from  3  to  10  inches.  They  are  the  most  abundant 
amphibian  in  the  Linton  beds.  The  present  genus  resembles  Lepterpeton  Huxley 
(334),  of  the  Kilkenny,  Ireland,  Coal  Measures.  But  that  genus  possesses  divided 
abdominal  rods,  or  "oat-shaped  scales,"  and  the  form  of  the  cranium  and  propor- 
tions of  the  body  are  different. 

The  genus  is  closely  related  to,  possibly  identical  with,  (Eslocep)ialus,  but  addi- 
tional material  will  be  required  to  settle  this  point. 

Cope  (123)  gives  the  following  key  for  the  separation  of  the  5  species: 

x.  Abdominal  rods  coarser,  not  more  than  10  in  5  mm. 

Median  pectoral  shield  discoid,  radiate  ridged;  muzzle  short P.  nummifer 

Median  pectoral  shield  oval,  pitted  and  ridged P.  marshii 

xx.  Abdominal  rods  hair-like,  15  or  more  in  5  mm. 

Median  pectoral  shield  with  radii  from  the  center,  the  principal  forming  a  cross;  form  wider. .   P.  vinchellianus 

Middle  pectoral  with  pits  at  the  center  and  few  or  no  radii;  form  narrow P.  pectinatus 

Middle  pectoral  shield  narrow,  closely  reticulate  medially,  and  radiate  towards  the  circumference;  size 

half  that  of  last P-  serrula 

Ptyonius  pectinatus  Cope. 

Cope,  Proc.  Acad.  Nat.  Sci.,  1868,  p.  216. 

Cope,  Trans.  Am.  Phil.  Soc,  xiv,  p.  20,  1869. 

Cope,  Trans.  Am.  Phil.  .Soc,  xv,  p.  266,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  377,  pi.  xxvii,  fig.  7;  xxviii,  figs.  2,  3,  6;  pi.  xxix,  fig.  2;  pi.  xxx,  fig.  2; 

pi.  xxxv,  figs.  1-3;  pi.  xli,  fig.  1,  1875. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  24,  pi.  8,  fig.  3,  1909. 
Schwarz,  Beitrage  zur  Paleontologie  und  Geologic  Osterreich-Ungarns  und  des  Orients,  Bd.  xxi,  p.  83, 

figs.  23,  24,  26,  1908. 

139 


140 


THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 


Type :  It  is  impossible  to  determine  which  one  of  the  specimens  is  the  type.  There 
are  numerous  representatives  of  the  species,  as  follows:  Nos.  140,  1096  G,  8345  G, 
8555  G,  1089  G,  2,  132,  133,  no  number,  1094  G,  8545  G,  8677  G,  1159  G,  105,  no 
number,  1091  G,  ya,  1092  G,  1093  G,  1095  G,  153,  and  others  unnumbered  in  the 
American  Museum  of  Natural  History;  in  the  U.  S.  National  Museum  are  the  fol- 
lowing: Nos.  4458,  4463,  4464,  4514.     (Plate  20,  fig.  2.) 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  most  abundant  species  of  the  Linton  Coal  Measures.  There  are  over  three 
dozen  specimens  preserved  in  the  Newberry  collection.  The  species  is  a  clearly 
marked  one,  as  a  rule,  though  there  is  great  variation  in  the  size  of  the  body  and  the 
form  of  the  vertebra?.  Though  there  are  several  apparently  complete  skulls  pre- 
served in  the  collection,  it  is  impossible  to  make  out  the  morphology  of  the  ele- 
ments on  account  of  the  amount  of  crushing  to  which  the  skulls  have  been  subjected. 

The  head  is  lancet-shaped,  and  the  muzzle  very  elongate,  slender,  and  acute  at 
the  extremity.  The  head  is  in  fact 
a  miniature  of  an  ichthyosaur 
cranium.  (Plate  20,  fig.  2.)  The 
orbits  are  large  and  posterior  to 
the  median  line.  The  anterior  por- 
tion of  the  skull  is  narrow,  poste- 
riorly truncate,  and  the  mandibular 
angle  is  projecting.  The  posterior 
portion  of  the  mandible  is  sculp- 
tured. Possibly  the  entire  cranium 
was  also,  and  this  has  been  lost; 
in  fact,  this  sculpturing  is  indicated 
in  one  or  two  specimens.  The  teeth 
are  conical  and  sharp,  longitudi- 
nally striate,  and  anisodont.  There  seemstobe  evidence  of  palatine  or  pterygoid  teeth, 
though  this  needs  confirmation.  The  pectoral  plates  are  well  preserved,  with  the 
interclavicle  a  narrow  oval,  with  anterior  and  posterior  prolongations.  In  one  speci- 
men it  is  sculptured.  The  clavicles  are  narrow  and  slightly  sculptured.  The  abdom- 
inal scutellae  are  bristle-like. 

The  vertebrae  are  short,  with  expanded  neural  and  haemal  spines.  The  expanded 
condition  of  the  neural  spines  begins  over  the  thoracic  region,  where  they  are  low. 
They  become  well  developed  in  the  posterior  dorsal  region.  The  caudal  fan-shaped 
spines  are  larger.  The  dilated  portions  form  equilateral  triangles  which  stand  on 
moderately  short  pedicels.  They  are  weakly  ridged,  and  each  ridge  is  prolonged  into 
a  narrow  acute  tooth  beyond  the  margin,  1 1  of  which  may  be  counted  on  one  of  the 
best -preserved  spines.  The  longitudinal  striae  are  terminated  near  the  pedicel  by 
two  others  which  cross  obliquely  from  each  side,  and,  meeting,  present  the  appear- 
ance of  the  margin  of  a  cup  sculptured  in  relief,  from  which  the  striae  arise.  Pedicels 
smooth.  The  spines  are  in  contact  at  their  angles,  thus  forming  a  continuous  line. 
In  a  typical  specimen  there  are  6  in  half  an  inch,  in  another  7,  and  in  a  third  8.  The 
ribs  are  well-developed  and  slender. 


Fig.  30. — Restoration  of  Ptyonius.    X  I. 


THE   AISTOPODOUS   MICKOSAURIAN    FAMILY    PTYONIID/E.  141 

No  traces  of  fore  limbs  have  been  detected  in  the  numerous  specimens,  but  ele- 
ments of  hind  limbs  are  preserved.  In  one  of  these  the  femur  is  a  small  bone, 
contracted  at  the  middle.    The  form  of  the  body  is  snake-like. 

There  were  probably  from  75  to  100  vertebrae  in  a  single  animal.  The  form  may 
be  well  compared  to  the  modern  Amphiuma  so  far  as  appearances  are  concerned; 
structurally  they  are  widely  separate.  This  species  is  one  which  is  peculiarly  char- 
acteristic of  the  Linton  fauna. 

Measurements  of  Ptvonius  i-ectinatus  Cope. 

Nos.  107  and  10Q4  C,  American  Museum.  Measurements  of  a  small  jaw,  No.  8555  C,  American 

Museum. 
mm.  mm. 

Length  of  specimen 137  Length  of  jaw 15 

Length  of  skull 26  Greatest  width 1.5 

Posterior  width  of  skull 8  Length  of  tooth I 

Intcrorbital  width 3  Measurements  of  specimen  No.  4458,  U.  S.  National 

1  Mameter  of  orbit 1.5  Museum. 

Vertical  expanse  of  vertebra 6  Length  of  specimen '. 65 

\\  ldth  of  neural  fan 2  Length  of  skull 22 

Diameter  of  pedicel I  Width  of  skull 6 

Ptyonius  vinchellianus  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  p.  177,  1871. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  376,  pi.  xxviii,  fig.  1,  1875. 

Type :  Specimen  in  the  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  is  represented  by  the  opposite  halves  of  a  single  specimen,  which 
includes  only  the  cranium  and  anterior  half  of  the  body.  The  fan-shaped  neural 
spines  commence  but  a  short  distance  behind  the  line  of  the  pectoral  shields.  They 
are  low,  with  a  few  coarse  ridges,  the  margin  being  entire.  The  abdominal  rods  are 
delicate  and  hair-like.  The  interclavicle  is  oval,  with  a  few  radiating  crests,  which 
originate  at  the  center;  in  the  areas  behind  there  are  a  few  scattered  tubercles. 
The  clavicles  are  ridged  near  the  margin. 

The  cranium  is  lanceolate  in  form,  and  the  bones  of  the  dorsum  are  marked  with 
a  few  raised  points  and  ridges.  The  species  is  about  the  size  of  Ptyonius  pectinuttis 
Cope,  and  differs,  apparently,  from  that  species  in  the  rather  insignificant  charac- 
ter of  a  narrower  interclavicle  and  in  the  ornamentation  of  the  same.  Dedicated  to 
Professor  Alexander  Winchcll,  of  the  University  of  Michigan. 

Measurements  of  Ptyonius  vinchellianus  Cope. 

mm. 

Length  of  cranium 20 

Width  of  same 8 

Length  of  interclavicle 4-2 

Ptyonius  marshii  Cope. 

Cope,  Trans.  Amer.  Phil.  Soc.,  xiv,  p.  24,  1869  (Colostcus  marshii). 

COPE,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  375,  pi.  xxvii,  fig.  6;  pi.  xxviii,  fig.  3,  1875. 

Cope,  Proc.  Am.  Phil.  Soc,  xn,  p.  177,  1871. 

Type:  Specimen  No.  1157  G,  American  Museum  of  Natural  History. 
Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  head  is  elongate  lanceolate.  The  upper  surface  of  the  frontal  bones  is 
punctate-rugose  in  relief,  with  short   radii  toward  the  margin.     The  distal  two- 


I42  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

thirds  of  the  mandible  is  narrow  wedge-shaped ;  the  external  surface  is  coarsely 
pitted.  There  are  no  teeth  preserved.  The  pectoral  elements  are  displaced,  but  the 
clavicles  are  subtriangular,  and  are  strongly  ridged  toward  the  inner  margin.  The 
interclavicle  is  short  spatulate,  the  narrow  portion  directed  anteriorly;  the  posterior 
rounded.  It  is  coarsely  pitted  medially,  and  coarsely  and  strongly  radiate-ridged  to 
the  margin.  The  abdominal  armature  commences  immediately  behind  the  pectoral 
girdle.  It  consists  of  elongate,  narrow,  subcylindric  scales,  which  meet  on  the 
median  line,  converging  anteriorly.    Small  limbs  are  present. 

Measurements  of  Ptyonius  marshii  Cope. 

Type  specimen:  mm. 

Length  of  fragmentary  skull 7 

Width  of  skull  as  preserved 5 

Length  of  entire  specimen 68 

Width  across  pectoral  plates 8 

Width  across  belly 7 

Specimen  No.  1098  G,  American  Museum: 

Length  of  specimen 32 

Width  of  specimen 7 

4  scutes  of  abdominal  armature  in  1  mm. 

Ptyonius  nummifer  Cope. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  pp.  374,  375,  pi.  xli,  figs.  2  and  3,  1875. 
Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  p.  356,  pi.  63,  fig.  3,  1909. 

Type:  Specimen  No.  8546  G,  American  Museum  of  Natural  History.  No. 
8614  G,  same  museum,  is  associated  with  the  type  specimen. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

Two  well-preserved  individuals  display  peculiarities  which  indicate  specific  dis- 
tinctness from  the  previously  known  species  of  Ptyonius.  The  abdominal  rods  are 
of  the  coarse  type  of  those  of  P.  marshii.  The  caudal  fans  are  well  developed,  and 
not  so  wide  as  in  P.  pectinatus.  The  interclavicle  is  a  discoid  body  of  different  form 
from  that  of  P.  marshii  and  I  can  not  detect  the  clavicles.  The  sculpture  consists 
of  strong  ridges,  which  radiate  from  the  center  to  near  the  border.  Immediately  in 
front  of  this  interclavicle  is  the  head,  which  has  a  different  form  from  that  of  the 
other  known  species.  The  interorbital  width  is  two-thirds  the  long  diameter  of  the 
orbit.  The  structure  of  the  skull  can  not  be  made  out.  A  slender,  elongate  hind 
limb  is  present  in  the  second  specimen,  and  a  humerus  is  well  preserved  in  the  type. 

Measurements  of  the  Type  of  Ptyonius  nummifer  Cope. 
(No.  8546  G,  American  Museum  of  Natural  History.    No.  8614  G  is  associated  in  the  same  species.) 

mm.  mm. 

Length  to  beginning  of  caudal  fans 65      Width  of  abdominal  armature 8 

Length  of  head 15      Length  of  a  caudal  fan 2.5 

Length  from  muzzle  to  orbits 6      Length  of  femur 5 

Length  of  interclavicle 7       Proximal  width  of  femur 1.5 

Width  of  interclavicle 8 

Ptyonius  serrula  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  1871,  p.  177. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  379,  pi.  xxviii,  fig.  5;  pi.  xxx,  fig.  I,  1875. 

Type:  Specimen  No.  8615  G,  American  Museum  of  Natural  History. 
Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  specimens  of  this  species  indicate  that  the  form  was  only  about  half  the  size 
of  Ptyonius  pectinatus.     The  interclavicle  is  narrower  and  more  reticulately  sculp- 


THE    AISTOPODOUS    MICROSAURIAN    FAMILY    PTYONIID^.  I43 

tured.    The  tail  is  relatively  longer.    Abdominal  rods  hair-like.    Ribs  distinct.    Small 
limbs  are  present  in  one  specimen. 

Measurements  of  Cope's  Type  of  Ptyonius  serrula. 

mm.  mm. 

Length  of  specimen 95  Width  of  clavicle 1.5 

Length  of  portion  of  skull  preserved 3  Length  of  vertebra I 

Length  of  interclavicle 4.5  Width  of  vertebra  from   tip  of  neural  spine  to  tip 

Width  of  interclavicle '. 2  of  haemal  spine 4 

Length  of  clavicle 4 

Another  specimen  (456  G,  American  Museum  of  Natural  History)  shows  some 
of  the  same  characters.  There  is  not  the  slightest  basis  for  the  support  of-  this  spe- 
cies, so  far  as  I  can  observe.  The  ones  mentioned  by  Cope  are  insufficient.  It  is  in 
all  probability  a  mutant  or  variety  of  Ptyonius  pectinatus. 

Genus  (ESTOCEPHALUS  Cope,  1868. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  218,  1868. 
Cope,  Trans.  Amer.  Phil.  Soc,  xiv,  16. 
Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  1868,  217. 
Cope,  Proc.  Amer.  Phil.  Soc,  1871,  41. 
Cope,  Geol.  Survey  Ohio,  n,  pt.  11,  380,  1875. 

Type :  (Estocephalus  remex  Cope. 

Form  slender  and  snake-like;  caudal  vertebras  with  elongated,  dilated,  sculp- 
tured neural  and  haemal  spines.  Cranium  lanceolate.  Teeth  numerous,  of  nearly 
equal  size.  No  pectoral  shields  known;  abdomen  protected  by  very  numerous 
bristle-like  rods,  which  converge  forwards.  A  pair  of  weak  posterior  limbs ;  branchi- 
hyal  bones  present. 

In  the  only  well-preserved  species  the  cranial  bones  exhibit  no  sculpture  from 
the  parietal  region  forward.  The  genus  is  not  very  distinct  from  Ptyonius,  but  it 
can  not  be  united  with  that  genus  until  more  complete  material  is  available.  The 
species  of  the  genus  share  with  Ceraterpeton,  Urocordylus  (334),  and  Ptyonius,  as 
well  as  Crossotelos  (98),  from  the  Permian  of  Oklahoma,  the  elongation,  sculpture, 
and  expansion  of  the  neural  and  haemal  spines.  There  are  but  2  species,  which 
Cope  distinguishes  by  the  following  characters: 

I.  Vertebras  elongate;  fan-like  caudal  processes  narrowed.    Size  large;  mandibular  teeth  of  unequal  lengths, 

with  the  apices  turned  backward (Estocephalus  remex 

II.  Species  only  known  from  cranial  bones  with  teeth.    Teeth  equal,  erect,  with  acute  conic  apices,  II  in 

5  mm (Estocephalus  reclidens 

(Estocephalus  remex  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  217,  1868  (Sauropleura  remex). 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  218,  1868  ((Estocephalus  amphiumianus). 

Cope,  Trans.  Amer.  Phil.  Soc,  xiv,  p.  17. 

Cope,  Geol.  Surv.  Ohio,  vol.  11,  pt.  11,  p.  381,  pi.  xxvii,  fig.  5;  pi.  xxxi,  fig.  1;  pi.  xxxii,  fig.  1;  pi.  xxxiii,  fig.  2; 

pi.  xxxiv,  fig.  4,  1875. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  27,  1909. 

Type:  Specimen  undetermined.  The  following  specimens  are  to  be  found: 
Specimens  of  GLstocephalus  remex  Cope  in  the  National  Museum,  Nos.  451 1,  4460, 
4478.  There  is  one  specimen  of  (Estocephalus  remex  in  the  University  of  Chicago. 
Specimens  of  the  species  in  the  American  Museum  of  Natural  History:  Nos.  121, 
8322  G,  8694  G,  no  number,  8656  G,  8583  G,  8659  G,  19,  120,  8655  G,  8662  G, 
8708  G,  8665  G,  112,  8663  G,  8581  G,  8658  G,  8660  G,  8700  G,  8469  G,  1102  G, 
1 152  G  142,  8381  G,  and  obverse,  21,  8664  G,  8672  G,  8592  G,  8684  G. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 


144 


THE    COAL    MEASURES    AMPHIBIA   OF    NORTH    AMERICA. 


This  species  is  one  of  the  most  abundant  of  the  Linton  Amphibia.  Cope  based 
his  description  of  the  species  on  9  specimens.  There  are  more  than  two  dozen  avail- 
able at  the  present  time,  the  majority  of  them  being  in  the  possession  of  the  Amer- 
ican Museum  of  Natural  History.  There  is  a  single  specimen  in  Walker  Museum 
of  the  University  of  Chicago  and  3  in  the  United  States  National  Museum.  The 
numbers  of  all  of  these  specimens  are  given  above.  The  material  consists,  for  the 
most  part,  of  fragmentary  portions  of  the  vertebral  column,  but  there  are  a  few 
skulls  more  or  less  complete,  though  none  are  sufficiently  well  preserved  for  a  com- 
plete analysis  of  the  characters.  The  specimens  indicate  an  animal  slightly  smaller 
than  the  modern  Amphiuma  means  of  the  Mississippi  River. 

It  will  not  be  necessary  here  to  enter  into  a  detailed  account  of  each  specimen, 
since  this  has  been  done  by  Cope,  and  a  careful  comparison  of  his  descriptions  with 
the  originals  indicates  that  his  observations  are  correct.  The  species,  as  suggested 
in  the  discussion  of  the  genus,  is  not  clearly  distinct  from  those  of  Ptyonius,  and  it 
has  largely  the  characters  of  that  genus.    The  cranium  is  long,  slender,  and  wedge- 


F10.  31. — Restoration  of  CEstoccphalus.     X  I. 

shaped.  The  teeth  are  numerous  both  in  the  maxillary  and  in  the  mandible,  one 
specimen  indicating  about  30  in  a  single  series.  They  are  all  uniformly  cylindrical, 
except  at  the  extremity,  where  they  are  flattened  and  expanded  so  as  to  produce  a 
longitudinal  edge,  which  is  carried  backward  on  a  recurvature  of  the  apex.  The 
bases  are  anchylosed  equally  and  without  enlargement,  and  no  part  of  the  shaft  is 
striate  or  grooved.  The  upper  surface  of  the  cranium  is  narrow,  with  the  median 
suture  distinct.    The  skull  surface,  with  that  of  the  mandible,  is  smooth. 

Characteristic  of  the  species  are  the  remarkable  length  and  slenderness  of  the 
fan-shaped  neural  and  haemal  spines,  and  the  absence  of  an  acute  serration  on  their 
margins.  In  this  species  the  spines  have  a  laminiform  expansion  at  the  base  in 
their  plane.  One  specimen  exhibits  the  pelvic  region,  including  a  portion  of  the 
tail.  The  ilium  has  an  expanded  anterior  extremity  and  is  directed  backwards 
and  somewhat  inwards  on  either  side  of  the  vertebral  column.  The  femur  is  nearly 
straight,  short,  contracted  medially,  and  expanded  distally.  The  tibia  is  shorter 
and  is  subcylindrical.  Beneath  the  ilium  the  last  chevron  of  the  abdominal  rods 
appears,  the  outer  extremities  rising  on  the  base  of  the  tail. 

The  pectoral  arch  is  almost  unknown,  and  Cope  based  the  distinction  of  Ptyo- 
nius and  CEstocephalus  on  the  absence  of  these  plates  in  the  latter  genus — an  uncer- 


THE   AISTOPODOUS   MICROSAURIAN   FAMILY   PTYONIIMs.  I45 

tain  characterization.  The  fore  limbs  are  indicated  by  a  humerus.  There  were 
possibly  from  75  to  100  vertebrae  in  the  entire  column.  The  animal  was  exclusively 
adapted  to  life  in  the  water  and  was,  without  doubt,  an  excellent  swimmer.  There 
are  preserved  in  one  specimen  portions  of  what  seem  to  be  hyobranchial  elements. 

Measurements  of  CEstocephalus  remex  Cope. 

mm.  mm_ 

Length  of  entire  caudal  series 195         Length  of  mandibular  dental  series 24 

Width  at  ninth  vertebra 21          Depth  of  mandible  at  middle ^ 

Width  at  thirty-sixth  vertebra 2          Nine  teeth  in 5 

Length  of  four  phalanges  in  place 14         Length  of  longer  teeth 2 

Expanse  of  fan  of  proximal  caudal 18.5      Length  of  first  hsmal  branchial 6 

Length  of  ilium 1 1.5      Length  of  sixth  vertebra  from  skull 7 

Length  of  femur 11. 5      Width  of  centrum 3 

Length  of  tibia 7 

CEstocephalus  rectidens  Cope. 

Cope,  Trans.  Amcr.  Phil.  Soc.,  p.  268,  Apr.,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  n,  p.  386,  pi.  xxvii,  fig.  3,  1875. 

Type:  Specimen  No.  9033,  American  Museum  of  Natural  History,  collection 
of  J.  S.  Newberry. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  is  indicated  by  a  left  dentary  bone,  with  its  teeth  and  external  sur- 
face preserved.  The  latter  is  nearly  smooth  and  without  sculpture.  The  outer 
face  is  convex,  and  the  general  form  is  slender,  but  not  curved  upward  at  the  extrem- 
ity. The  extremity  of  the  dentary  does  not  show  any  evidences  of  teeth.  Teeth 
straight  and  conic,  apex  acute,  non-plicated. 

Cope  also  associated  with  this  species  a  portion  of  a  caudal  series,  consisting 
of  25  vertebrae.  The  centra  are  elongate  and  expanded  at  the  extremities.  The 
neural  arches  have  a  close  union.  The  neural  and  haemal  spines  are  fan-shaped  and 
striated.     The  bases  are  quite  narrow. 

Measurements  of  QSstocephalus  rectidens  Cope. 

mm.  mm. 

Length  of  dentary 22  Length  of  3  centra  of  the  caudal  series 8.6 

Length  of  tooth  line 15.2  Extent  of  neural  and  haimal  spines 8.7 

Depth  of  dentary  at  last  tooth 2.7 

Genus  THYRSIDIUM  Cope,  1875. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  365,  pi.  xxxi,  fig.  2,  1875. 

Type:  Thyrsidium  fascicular  e  Cope. 

Established  on  a  species  which  presents  its  principal  peculiarities  in  the  struc- 
ture of  the  vertebrae.  Two  specimens  present  inferior  views  of  the  spinal  column, 
showing  that  the  genus  possesses,  like  Siren,  enlarged  diapophyses,  but  they  are 
peculiar  in  their  fan-like  form.  They  resemble  slightly  the  neural  spines  of  the 
caudals  of  Ptyonius,  but  are  present  on  the  dorsal  vertebrae.  Whether  the  caudals 
of  the  present  species  possess  ornamented  neural  spines  the  specimens  do  not  indi- 
cate. The  abdomen  is  protected  by  the  usual  hair-like  rods  arranged  en  chevron, 
the  angle  directed  forwards.  No  indications  of  limbs  can  be  discovered  on  the 
blocks. 


I46  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

Without  the  cranial  bones  the  affinities  of  this  genus  can  not  be  determined; 
while  it  may  be  allied  to  Cocytinus,  the  vertebrae  of  that  form  are  without  peculiar 
diapophyses. 

Thyrsidium  fasciculare  Cope. 
Cope,  Geol.  Surv.  Ohio,  n,  pt.  n,  p.  365,  pi.  xxxi,  fig.  2,  1875. 

Type:  Specimen  No.  8552  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  best  preserved  example  of  this  species  includes  9  vertebrae  and  the  corre- 
sponding ventral  armature.  The  centra,  seen  from  below,  are  much  contracted  in 
their  form,  presenting  an  obtuse  median  rib,  which  expands  to  the  articular  extrem- 
ities. In  one  or  two  instances  the  latter  are  divided  by  fracture,  and  the  moder- 
ately concave  form  of  the  adjacent  surfaces  is  displayed.  The  diapophyses  are  of 
complex  form,  but  the  details  are  concealed  by  the  prevalent  thin  layer  of  coal  which 
invests  them.  An  inferior  prominence  runs  parallel  to  the  centrum;  outside  of  this 
the  process  is  obscurely  trilobate  and  thickened,  not  flattened,  as  in  the  caudal  ver- 
tebras of  Ptyonius.  Several  ribs  (fig.  8)  of  moderate  thickness  appear  by  the  side 
of  the  diapophyses.    Eleven  abdominal  ribs  in  5  mm. 

The  second  specimen  was  originally  referred  to  QLstocephalus  rcmcx,  as  a  pos- 
terior portion  of  its  vertebral  column,  immediately  preceding  the  caudal  series. 

This  reference  appears  to  be  incorrect,  although  the  resemblance  between  the 
corresponding  parts  in  the  two  genera  is  no  doubt  considerable,  and  the  alternative 
of  proposing  a  new  genus  and  species  was  not  at  that  time  advisable. 

The  neural  spines  are  longer  than  high,  and  are  nearly  in  contact  at  their  mar- 
gins; each  is  marked  by  about  5  obtuse  vertical  ribs.  A  fractured  section  of  the 
abdominal  spines  in  place  displayed  at  least  six  layers  of  them. 

The  material  on  which  the  above  account  is  based  is  imperfect.  The  spec- 
imen figured  by  Cope  (Geol.  Surv.  Ohio,  vol.  11,  pt,  11,  pi.  xxxi,  fig.  2,  1875)  is 
undoubtedly  a  portion  of  the  vertebral  column  of  QLstocephalus  remex.  Nos.  4462 
and  4480  of  the  United  States  National  Museum  may  be  representatives  of  Thyr- 
sidium fasciculare,  but  they  are  more  probably  QLstocephalus  remex;  if  they  are  the 
latter  this  leaves  the  type  as  the  only  known  specimen  of  the  species. 

Measurements  of  Type  of  Thyrsidium  fasciculare  Cope. 


mm. 


Length  of  portion  of  vertebral  column  preserved 70 

Width  of  ventral  armature 18 

Length  of  a  vertebra 9 

Width  of  same  vertebra 4 


CHAPTER  XX. 

THE  MICROSAURIAN  FAMILY  MOLGOPH1D/E,  FROM  THE  COAL  MEASURES  OF  OHIO 

AND  MAZON  CREEK,  ILLINOIS. 

Family  MOLGOPHID^  Cope,  1875. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  357,  1875. 
Cope,  Bull.  U.  S.  Nat.  Mus.,  No.  1,  p.  11,  1875. 

Type  of  family :  Molgophis. 

Body  long,  serpentine,  a  few  species  apparently  limbless,  ribless,  and  with 
abdominal  armature  lacking.  Vertebras  elongate,  neural  and  haemal  spines  short 
or  absent.  Ribs  long,  heavy,  and  broad.  The  vertebrae  seem  to  bear  the  charac- 
teristic marks  of  the  family.  One  species  has  the  skeleton  reduced  to  a  lanceolate 
skull  and  a  string  of  about  50  slender  vertebras,  all  the  rest  of  the  skeleton  being 
absent.  The  family  is  very  poorly  known,  but  was  apparently  of  wide  distribu- 
tion in  North  America  and  confined  to  this  continent.  The  representatives  of 
the  group  are  known  from  Iowa,  Illinois,  and  Ohio. 

Four  genera  are  assigned  to  the  family,  but  future  discoveries  will  undoubtedly 
demand  revision  of  the  present  classification.  The  genera  are  Molgophis,  Pleu- 
roptyx,  Phlegethontia,  and  Erpetobrachium.  The  distinguishing  characters  of  these 
genera  are  apparent  from  the  descriptions  of  the  various  forms.  The  skeletons  of 
the  species  are  too  incompletely  known  to  allow  the  establishment  of  a  tabular  key 
to  the  genera. 

Genus  MOLGOPHIS  Cope,  1868. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  220,  1868. 
Cope,  Trans.  Amer.  Phil.  Soc,  xiv,  p.  20,  1869. 
Cope,  Geol.  Surv.  Ohio,  it,  pt.  11,  p.  368,  1875. 
Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  263,  1874. 

Type:  Molgophis  macrurus  Cope. 
Cope  (123)  gives  the  following: 

"The  characters  of  this  genus  are:  body  long,  serpentine,  without  dermal  armature, 
so  far  as  known;  vertebras  long  and  broad,  with  very  prominent  zygapophyses  and  moder- 
ate neural  spines;  ribs  large,  curved.  No  limbs  or  cranium  can  be  ascribed  to  the  type  of 
the  genus.  The  ribs  are  long,  and  though  the  head  is  not  bifurcate,  there  appears  to  be 
both  tubercle  and  head  on  the  dilated  extremity.  Where  crushed  they  display  a  large 
median  vacuity. 

"This  genus  differs  from  Ophiderpeton  Huxley  (334)  in  the  characters  of  the  dorsal 
vertebrae,  which,  in  their  projecting  zygapophyses,  resemble  those  of  Amphiuma.  The 
lack  of  ventral  armature  distinguishes  it  from  CEstocephalus,  while  its  well-developed  ribs 
separate  it  from  Phlegethontia.'" 

Molgophis  macrurus  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  220,  1868. 

Wyman,  Am.  Jour.  Sci.  and  Arts,  p.  II,  fig.  I,  1858  (refers  to  a  batrachian  reptile). 

Cope,  Trans.  Amer.  Phil.  Soc,  xiv,  p.  20,  1869. 

Cope,  Trans.  Amer.  Phil.  Soc,  xv,  p.  263,  1874. 

Cope,  Geol.  Sun'.  Ohio,  11,  pt.  U,  p.  368,  pi.  xliii,  fig.  I,  1875. 

Type:  Specimen  No.  8617  G,  American  Museum  of  Natural  History,  collec- 
tion of  Dr.  J.  S.  Newberry. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

147 


148 


THE   COAL   MEASURES    AMPHIBIA   OF    NORTH    AMERICA. 


This  species  is  established  by  remains  of  2  individuals,  one  embracing  16  and 
the  other  14  vertebrae,  with  ribs.  The  neural  arches,  viewed  from  above,  have  a 
V-shaped  outline  posteriorly,  from  the  fact  that  the  broad  zygapophyses  meet  on  the 
median  line  and  spread  out  distally  over  the  broad  anterior  ones  adjoining.  The 
latter  appear  to  be  somewhat  concave  and  to  border  the  former  exteriorly  as  well  as 
inferiorly.  The  base  of  the  neural  spine  extends  to  the  posterior  emargination, 
but  not  quite  to  the  anterior.  The  breadth  of  the  dorsal  vertebrae  above  is  equal 
from  the  emargination  behind  to  the  anterior  margin  of  the  anterior  zygapophyses. 

The  ribs  are  long  for  an  amphibian,  but  not  long  for  a  reptile.  They  are  well 
curved,  chiefly  near  the  proximal  extremity.  The  longest  found,  measured  by  a 
cord,  equals  two  and  two-fifths  vertebrae.  These  vertebrae,  measured  along  the 
median  line  above,  equal  11  lines;  one  of  these  is  3.6  lines  in  width  above.  This 
animal  has  been,  like  Amphiama,  a  snake-like  amphibian,  but 
was  probably  still  larger.  How  near  the  affinities  to  this 
genus  may  be  can  not  now  be  determined,  owing  to  the  want 
of  many  important  parts  of  the  skeleton,  but  it  differs  in  the 
important  feature  of  large,  well -developed  ribs.  The  size  of 
the  vertebrae  would  indicate  a  body  of  the  size  of  the  com- 
mon rattlesnake  (Crotalus  horridus)  and  too  large  for  Br achy  - 
dectes  newberryi,  which  is  only  known  from  jaws. 


Molgophis  brevicostatus  Cope. 
Cope,  Geol.  Surv.  Ohio,  n,  pt.  11,  p.  369,  pi.  xliv,  fig.  1,  1875. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  27,  1909. 


Neura, 
spine 


Ribj 


Centrum 


Type:  Specimen  No.  8341  G,  Amer.  Mus.  of  Nat.  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

Represented  by  portions  of  the  vertebral  column  of  several 
individuals.  One  of  these  includes  9  pairs  of  ribs,  with  verte- 
brae, and  another  13  pairs.  The  vertebrae  are  subquadrate  in 
section,  and  the  concavity  of  the  two  articular  faces  is  not 
deep.  They  support  strong  lateral  ridges  separated  by  deep 
concavities.  The  heads  of  the  ribs  are  somewhat  contracted, 
and  the  shafts  present  outward  a  tubercular  angle  at  a  dis- 
tance of  one-fourth  the  length  from  the  head.  The  remaining 
part  of  the  shaft  is  stout,  nearly  straight,  and  gradually  con- 
tracts to  an  obtuse  extremity.  It  possesses  a  narrow  medullary  cavity.  In  none 
of  the  specimens  is  there  any  trace  of  abdominal  armature,  but  abundant  remains 
of  the  contents  of  the  abdominal  cavity,  in  proper  position,  are  preserved  on  the 
blocks.  This  species  is  more  massive  than  Molgophis  macrurus,  and  the  ribs  are 
shorter,  thicker,  and  less  curved. 

Ventral  scutellae  are  present  in  one  specimen  of  this  species.  There  are  a  number 
of  specimens.  They  have  the  following  numbers  at  the  American  Museum:  158, 
1 100  G,  no  number,  8341  G  (type),  no  number,  8466  G;  and  4477  in  the  U.  S. 
National  Museum. 


Fig.  32. — Drawing  from 
Cope's  figure  of  Molgo- 
phis brevicostatus  Cope. 
X0.5. 


THE   MICROSAURIAN    FAMILY   MOLGOPHID^E.  I49 

Measurements  of  Type  Specimen  of  Molgophis  brevicostatus  Cope. 

mm.  mm. 

Length  of  7  vertebra 105  Length  of  a  rib  on  a  curve 24 

Length  of  1  centrum 16  Greatest  thickness  of  same 2.5 

Diameter  of  same  vertically 11 

Molgophis  wheatleyi  Cope. 

Cope,  Gcol.  Surv.  Ohio,  II,  pt.  n,  pp.  369,  370,  pi.  xlv,  fig.  1,  1875. 
Cope,  Trans  Am.  Phil.  Soc,  xv,  p.  263,  1874. 

Type:  Specimen  No.  1101  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

A  critical  study  of  the  type  specimen  of  this  species  does  not  reveal  anything 
essentially  different  from  the  description  of  Cope.  The  following  is  taken  from  his 
report (123): 

"Established  on  a  specimen  which  exhibits  about  twenty-five  vertebrae  with  ribs,  and 
the  posterior  portion  of  the  cranium.  No  traces  of  abdominal  scales  or  rods,  thoracic 
shields,  or  limbs  are  visible.  By  such  negative  characters  it  is  referable  to  the  genus  Mol- 
gophis, although  the  definition  of  this  genus  is  incomplete.  The  present  batrachian  may, 
indeed,  be  ultimately  found  to  be  an  Ophiderpeton,  to  which  it  also  bears  some  resemblance. 

"The  specimen  is  that  of  an  animal  of  very  much  smaller  size  than  the  M.  macrurus. 
The  vertebra  are  of  moderate  length,  with  a  low  neural  spine,  and  centrum  angular  at 
the  sides  and  truncate  at  the  articular  extremities  when  in  place.  The  ribs  are  rather 
short,  slightly  curved,  apparently  hollow  and  intercentral  in  position.  Although  the  ver- 
tebral centra  are  ossified,  the  elements  of  the  cranium  have  a  larval  appearance.  These 
consist  of  two  parallel  bony  plates,  which  resemble  the  fronto-parietal  bones  of  the  frog; 
they  are  slightly  separated  from  each  other,  but  do  not  inclose  a  fontanelle.  A  wedge- 
shaped  bone  extends  from  the  outside  of  the  front  of  each  of  these,  acuminate  behind, 
and  widening  anteriorly  in  the  position  of  a  postfrontal  bone.  In  front  of  the  posterior 
border  of  each  parietal,  on  its  outer  side,  a  bony  enlargement  arises  which  contracts  out- 
ward and  forward  into  a  narrow  element  which  curves  forward  beneath  the  postfrontal. 
These  look  like  an  anteriorly  directed  quadrate  with  articular  bone,  such  as  seen  in  the 
larvae  and  some  adults  of  existing  batrachians.  These  determinations  will  require  con- 
firmation from  additional  material.  In  the  meantime  it  is  evident  that  the  present  specimen 
can  not  be  referred  to  any  of  the  other  species  herein  described.  The  elements  of  the 
cranium  are  entirely  smooth  with  no  sign  of  sculpture,  and  in  this  respect  the  present 
species  is  unlike  any  of  the  other  known  from  the  Carboniferous." 

The  vertebrae  are  not  so  clearly  marked  as  one  is  led  to  believe  from  Cope's 
figure. 

Measurements  of  the  Type  of  Molgophis  wheatleyi  Cope. 

mm.  mm. 

Length  of  entire  specimen 63  Width  of  a  vertebra. . 1.5 

Length  of  portion  of  skull  preserved 9  Length  of  a  posterior  rib 5 

Posterior  width  of  same 8.5  Width  of  rib 5 

Length  of  a  vertebra 1.5 

The  species  is  dedicated  to  Charles  M.  Wheatley,  of  Phcenixville,  Pennsylvania, 
one  of  the  original  investigators  of  the  Linton  deposits.  It  is  a  part  of  the  New- 
berry Collection. 

Additional  material  of  this  species  is  represented  by  specimens  Nos.  7  and  8699  G 
of  the  American  Museum  of  Natural  History.  They  are  both  very  unsatisfactory. 
They  consist  of  molds  of  the  vertebral  column,  with  in  one  case  an  enlargement  at 
one  end  which  may  represent  the  head,  and  if  such,  the  specimen  probably  repre- 
sents a  distinct  species.     The  impression,  No.  7,  contains  molds  of  about  30  ver- 


150  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

tebrae  which  are  very  similar  in  form  to  those  exhibited  by  the  type  of  the  species. 
To  the  vertebrae  are  articulated  short,  curved  ribs  of  a  slender  nature.  The  ver- 
tebras themselves  are  short  and  somewhat  constricted  in  the  middle. 

The  other  impression,  8699  G,  contains  impressions  of  about  20  vertebrae,  appar- 
ently immature,  though  one  can  not  be  entirely  sure  as  to  the  nature  of  the  struc- 
tures. They  are  covered  over  with  a  thin  layer  of  carbonaceous  material  which  is 
impossible  to  remove  satisfactorily.  The  two  specimens  remind  one  of  what  Hux- 
ley has  written  in  regard  to  the  forms  of  Microsauria  (334)  from  Kilkenny,  Ireland. 

Measurements  of  Nos.  7  and  8699  G  (Molgophis  whf.atleyi). 

mm.  mm. 

Length  of  No.  7 87       Length  of  vertebra 2 

Length  of  head  mold 18       Length  of  rib 3 

Posterior  width  of  head 6       Length  of  specimen  No.  8699  G S3 

Genus  ERPETOBRACHIUM   Moodie,   1912. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  353,  1912. 

Type:  Erpetobrachium  mazonensis  Moodie. 

The  generic  characters  are  apparent  in  the  greatly  elongated  fore  limb,  in  the 
exceptionally  broad  scapula,  the  long  radius  and  ulna,  which  slightly  exceed  the 
humerus  in  length,  a  character  hitherto  unknown  among  Carboniferous  Amphibia. 

Erpetobrachium  mazonensis  Moodie. 
Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  pp.  353-354,  pi.  2,  fig.  2;  pi.  8,  fig.  3,  1912. 

Type:  Specimen  No.  799  (222),  Yale  University  Museum. 

Horizon  and  locality :  Mazon  Creek  shales,  near  Morris,  Illinois. 

The  scapula  of  the  present  form  is  exceptional  in  its  shape.  It  resembles  an 
asymmetrical  pyramid,  the  anterior  side  of  the  lower  edge  of  the  bone  being  con- 
tracted so  that  the  anterior  edge  of  the  element  is  arcuate.  Its  top  is  very  thin  and 
possibly  terminated  in  a  broad  cartilage.  The  lower  end  is  thick  and  heavy  and 
the  articular  surface  is,  apparently,  well  formed,  though  somewhat  obscured. 

The  element  identified  as  a  clavicle  is  lying  on  its  edge  and  has  the  proportions 
of  the  clavicle  of  Mazonerpeton  costatum.  The  exterior  edge  is  somewhat  rounded 
and  small.  A  portion  of  another  element  which  I  suppose  to  represent  the  coracoid 
lies  alongside  the  humerus,  although  its  form  is  quite  obscured.     (Plate  3,  fig.  3.) 

The  humerus  has  a  remarkably  well-formed  head.  In  perfection  of  formation 
it  corresponds  well  with  that  of  the  higher  reptiles.  This  surface  can  even  be  divided 
into  an  anterior  and  a  posterior  articulation.  The  element  projects  posteriorly  for 
the  distance  of  1  mm.  from  the  surface  of  the  shaft.  The  shaft  immediately  below 
the  head  is  somewhat  flattened  and  has  an  ovoid  section.  Further  on  it  becomes 
flattened,  a  part  of  which  is  probably  due  to  pressure  during  fossilization. 

The  elements  of  the  forearm  are  both  preserved  and  are  approximately  equal  in 
size.  They  are  remarkable  in  that  they  exceed  the  humerus  in  length,  although 
they  are  not  so  heavy  as  that  element.  They  are  greatly  elongate  and  slender,  with 
the  middle  of  the  shaft  only  moderately  contracted.  The  articular  surfaces  are 
well  formed  and  both  bones  were  hollow,  as  was  also,  apparently,  the  humerus. 
The  ulna  may  be  represented  by  the  most  posterior  of  the  two  elements,  though 
the  relations  of  the  elements  may  have  been  reversed  (fig.  15,  D). 


THE   MICROSAURIAN   FAMILY  MOLGOPHID^.  151 

The  base  of  the  left  wing  of  an  orthopterous  insect  possibly  allied  to  Paolia 
gitrlcyi  Scudder  lies  between  the  radius  and  ulna.  The  nodule  also  contains  impres- 
sions of  plants,  a  portion  of  a  frond  of  a  Neuropteris  and  the  impression  of  one  of 
the  Cordaites.  Lying  next  the  radius  is  a  slender  elongate  element  which  may  be  a 
rib  or  a  portion  of  a  metacarpal.  If  a  rib,  it  indicates  that  the  animal  belongs  among 
the  Branchiosauria.  The  fragment  is  only  half  as  long  as  the  radius  and  is  entirely 
too  obscure  to  base  any  conclusions.  The  other  characters  of  the  specimen  point 
quite  strongly  to  its  microsaurian  affinities. 

The  structure  of  the  articular  surfaces  of  the  limb  bones  alone  would  indicate 
the  microsaurian  relationship  of  Erpetobrachium.  It  may  be  provisionally  asso- 
ciated in  the  family  Molgophidae  with  such  forms  as  Molgophis  brevicostatus  Cope, 
Molgophis  {Pleuroptyx)  clavatus  Cope,  and  Molgophis  macrurus  Cope  from  the  Coal 
Measures  of  Linton,  Ohio. 

Measurements  of  the  Type. 

mm.  mm. 

Length  of  scapula 14      Diameter  of  shaft 2 

Distal  width 6       Distal  width 3 

Proximal  diameter 3      Length  of  radius 25 

Length  of  clavicle  (?) 24      Proximal  width 4 

Length  of  humerus 25       Diameter  of  shaft 3 

Length  of  ulna 24      Width  of  distal  end 4 

Proximal  width 4 

Genus  PLEUROPTYX  Cope,   1875. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  370,  pi.  xlii,  fig.  1;  pi.  xliv,  fig.  2,  1875. 
Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  16,  1875. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  27,  1909. 

Type:  Pleuroptyx  clavatus  Cope. 

The  specimens  on  which  the  species  of  this  genus  repose  do  not  exhibit  crania. 
The  5,  probably  6,  specimens  which  represent  them  offer  various  views  of  the  ver- 
tebral column,  and  in  none  is  there  any  trace  of  ventral  or  thoracic  armature.  Limbs 
can  be  ascribed  to  them  with  probability  only.  The  vertebrae  are  of  moderate 
length,  with  well-developed  zygapophyses,  and  a  short  and  not  very  elevated  neu- 
ral spine  in  the  dorsal  region,  which  is  not  sculptured  in  any  way.  The  generic 
character  is  seen  in  the  ribs.  These  are  rather  short  and  very  stout  and  support  an 
ala  on  the  posterior  or  convex  border,  which  expands  downwards,  and  then  sud- 
denly contracts  to  the  shaft.  The  extremity  of  the  latter  is  broad  and  truncate, 
and  includes  a  medullary  cavity,  which  is  only  partially  fitted  with  cancellated 
tissue. 

Pleuroptyx  clavatus  Cope. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  370,  pi.  xlii,  fig.  1 ;  pi.  xliv,  fig.  2,  1875. 
Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  16.  1875, 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  27,  1909. 

Type:  Specimen  No.  8617  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  general  appearance  of  the  species  of  Pleuroptyx  is  that  of  the  Molgophis,  so 
far  as  known,  but  nothing  resembling  the  peculiar  structure  of  the  ribs  is  seen  in 
any  other.    There  is  no  assurance  that  the  genus  is  distinct  from  Molgophis. 


152 


THE    COAL   MEASURES    AMPHIBIA    OF    NORTH    AMERICA. 


Humerus 


Ulna. 


Parts  of  two  individuals  express  the  typical  characters  of  this  species,  while  a 
third  only  differs  in  being  considerably  smaller.  A  fourth  may  very  probably  be 
referred  here,  and  another,  bearing  several  elements  of  a  leg,  should  be  most  likely 
associated  with  the  last  mentioned. 

The  ribs  are  considerably  narrowed  near  the  head,  and  appear  to  possess  a  low 
tubercular  process  some  distance  below  it.  The  shaft  is  curved 
throughout;  the  laminar  expansion  is  quite  thin;  while  the  distal 
end  is  expanded  and  concave,  perhaps  for  the  attachment  of  car- 
tilage, although  no  trace  of  this  remains  on  the  shale.  The  neural 
spines  have  short  bases,  oblique  anterior  and  nearly  straight  pos- 
terior borders,  with  obtuse  extremity.  I  perceive  no  essential  dif- 
ference in  the  smaller  specimen,  which  is  one-third  less  than  the 
types. 

The  limb  is  appropriate  in  its  proportions  to  the  present 
species,  and  may  be  described  in  this  place.  The  first  segment 
is  one-third  longer  than  the  second,  and  has  a  transversely 
expanded  head.  The  shaft  is  stout,  the  distal  extremity  not 
expanded  and  concave.  The  second  segment  is  stout,  more  ex- 
panded proximally  than  distally,  the  proximal  end  truncate  and 
slightly  concave.  A  bone,  much  displaced,  lies  near  it,  and  is  fig.  33. 
probably  ulna  or  radius;  it  is  as  stout  as  the  first,  the  end  not 
expanded.  Of  metatarsals  there  are  2,  three-fifths  the  length 
of  the  second  bone  of  the  leg,  and  of  phalanges  2,  of  2  digits 
each.  The  proximal  are  three-fourths  the  length  of  the  metatarsals,  and  indicate 
elongate  toes.  The  obverse  of  the  specimen  is  preserved,  and  contains  no  addi- 
tional toes  or  phalanges. 


Phalanges 


Fore  limb  of  a 
member  of  the  Mol- 
gophida-,  possibly 
Pleuroptyx  clavatus 
Cope.     X0.75. 


Measurements  of  the  Type  of  Pleuroptyx  CLAVATUS  Cope. 


Length  of  vertebral  centrum 14 

Depth  of  a  vertebral  centrum 9 

Depth  of  entire  vertebra 22 

Length  of  neural  spine 8 

Height  of  a  neural  spine 6 

Length  of  a  rib  on  the  curve 42 


Width  of  a  rib  at  ala 9 

Width  of  a  rib  at  extremity 5 

Length  of  first  segment  of  leg 38 

Length  of  second  segment  of  leg 24 

Length  of  metapodial  bone 10 

Length  of  first  phalanx 7 


Hitherto  only  two  portions  of  the  dorsal  series  and  a  left  limb  have  been  assigned 
to  this  species.  The  present  specimen  (No.  4479,  U.  S.  Nat.  Mus.)  thus  proves  of 
interest  in  determining  that  the  creature  was  long-tailed,  like  CEstocephaius,  Ptyo- 
nius,  and  Phlegethontia,  but  unlike  the  first  two  genera  the  neural  and  haemal  spines 
are  not  elongate  nor  marked  with  radiating  lines.  The  neural  spines  are  indistinct 
and  if  developed  at  all  were  very  low  and  short. 

The  centra  are  short,  cylindrical,  and  thick.  They  gradually  decrease  in  size 
to  where  they  are  lost,  since  the  portion  preserved  does  not  represent  the  entire 
length  of  the  tail.  There  may  have  been  15  more  vertebrae  distally  and  5  more 
proximally,  thus  making  about  75  caudal  vertebrae,  as  Woodward  (630)  has  deter- 
mined obtains  in  Ceraterpeton  galvani  Huxley. 


THE    MICROSAURIAN    FAMILY   MOLGOPHID^.  153 

The  ribs  are  continuous  throughout  the  length  of  the  tail  preserved  and  have 
precisely  the  same  structure  as  is  found  in  the  dorsal  region  with  the  possible  excep- 
tion that  the  posterior  alar  expansion  is  not  so  well  developed  in  the  caudal  ribs. 
The  ribs  are  decidedly  fan-shaped  and  articulate  by  a  single  head  with  a  short 
transverse  process.    They  are  distinctly  curved  like  all  microsaurian  ribs. 

Measurements  of  Specimen  of  Pleuroptyx  clavatus  Cope. 

(No.  4509,  U.  S.  National  Museum,  Linton,  Ohio,  Coal  Measures.) 

mm.  mm. 

Length  of  tail  as  preserved 105  Length  of  rib 5 

Length  of  anterior  vertebra 1.5  Width  of  rib I 

Diameter  of  anterior  vertebra 1 

The  above-described  specimen  represents  what  I  suppose  to  be  the  posterior 
part  of  the  body  of  Pleuroptyx  clavatus  Cope.  The  characters  of  the  ribs  and  ver- 
tebras are  the  same.  The  fragment  is  interesting,  since  it  gives  an  insight  into  the 
form  of  the  body,  which  was  slender,  conforming  thus  to  other  long-tailed  micro- 
saurs.    Length  of  specimen,  65  mm. ;  width  of  specimen,  30  mm. 

There  is  still  a  third  example  of  this  species  among  the  collections  belonging  to 
the  National  Museum  (No.  4484).  The  specimen  includes  a  badly  crushed  pos- 
terior portion  of  a  skull  and  a  series  of  about  16  crushed  vertebrae,  with  several 
pairs  of  ribs  and  ventral  scutes. 

Very  little  can  be  said  of  the  skull  save  that  the  maxilla  of  the  right  side  was 
long  and  bore  from  15  to  20  teeth,  of  which  9  are  preserved  more  or  less  completely. 
The  mandible  is  likewise  crushed  and  one  can  not  determine  its  elements.  Por- 
tions of  2  or  3  teeth  are  preserved.    The  form  of  the   mandible  is  long  and  slender. 

The  ventral  scutes  are  of  the  pectoral  region.  They  are  long,  slender,  and  thread- 
like.   They  are  not  closely  packed,  but  I  count  12  in  a  distance  of  a  millimeter. 

So  far  as  can  be  determined  the  vertebrae  are  the  same  as  has  been  described 
for  other  specimens.  They  are  short  and  heavy.  The  ribs  show,  for  the  most  part, 
the  same  characters  as  the  type  specimen. 

Measurements  of  Third  Specimen  (No.  4484,  U.  S.  Nat.  Mus.). 

mm.  mm. 

Length  of  specimen 60  Length  of  anterior  vertebra 2 

Length  of  preserved  portion  of  skull 15  Length  of  rib 4 

Length  of  tooth 25  Width  of  rib I 

Posterior  width  of  mandible 3 

Genus  PHLEGETHONTIA  Cope,   1871. 

Cope,  Proc.  Am.  Phil.  Soc.,  p.  177,  1871. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  366,  1875. 

Type:  Phlegethontia  linearis  Cope. 

This  is  one  of  the  most  interesting  genera  of  the  present  series.  It  rests  chiefly 
on  a  single  specimen  of  one  species,  which  is  not  perfect,  but  which  displays  the 
following  characters:  Head  elongate-triangular;  body  and  tail  extremely  elongate, 
the  dorsal  vertebrae  without  ribs,  and  the  caudals  without  dilated  spines ;  no  ven- 
tral armature  nor  limbs.  As  a  great  portion  of  the  length  is  presented,  and  no  ventral 
rods  or  scales  are  visible,  and  as  this  character  is  confirmed  by  a  second  specimen, 
it  probably  belongs  to  the  genus.    The  pectoral  shields  are  also  wanting  in  the  spec- 


154  THE    COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

imen,  but  as  there  is  a  considerable  vacuity  behind  the  skull  of  the  specimen,  it 
may  be  that  these  were  lost  with  other  parts.  Chevron  bones  are  not  observable 
on  the  caudal  vertebrae.    This  form  is  a  true  amphibian  snake. 

Phlegethontia  linearis  Cope. 
Cope,  Geol.  Surv.  Ohio,  II,  pt.  H,  p.  367,  pi.  xliii,  fig.  2,  1875. 

Type:  Specimen  in  the  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

In  the  only  specimen  the  dorsal  vertebrae  are  much  involved  anteriorly,  so  that 
the  length  is  not  readily  ascertained.  There  is  an  outline  of  a  triangular  object 
which  may  represent  the  skull  of  this  specimen,  although  it  is  so  far  removed  from 
the  vertebrae  that  there  is  some  doubt  as  to  whether  it  belongs  with  the  vertebrae 
or  not.  Indeed,  there  is  even  doubt  whether  it  is  a  skull.  The  vertebrae  have 
longitudinal  diapophysial  keels,  and  have  a  zig-zag  interlocking  of  neural  arches. 
The  latter  are  distinctly  turned  outward.  The  vertebrae  are  very  numerous,  and 
the  tail  very  attenuated.  The  number  preserved  is  about  60.  The  total  length  of 
the  coils  unwound  is  about  295  mm.,  or  11  coils  in  8  lines;  but  there  are  interrup- 
tions not  measured  and  confusions  not  unraveled. 

This  is  the  most  elongate  and  slender  of  all  the  species  of  the  Carboniferous 
Amphibia.  The  vertebrae  are  apparently  ribless  and  there  are  no  evidences  of 
limbs  or  pectoral  plates.  It  may  be  said  that  the  body  consists  entirely  of  skull 
and  vertebrae. 

Measurements  of  the  Type  of  Phlegethontia  linearis  Cope. 

Entire  length  of  skull  (?) 18  mm.  Length  of  single  vertebra 2.50  mm. 

Width  of  same 8  mm.  Height  of  vertebra 1 .50  mm. 

Length  of  vertebral  column  as  preserved. . .     295  mm.  Estimated  length  of  body 15        in. 

No.  8370  G,  American  Museum  of  Natural  History,  shows  a  few  vertebrae. 

Phlegethontia  serpens  Cope. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  367,  pi.  32,  fig.  2,  1875. 

Type:  Specimen  No.  1102  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  amphibian  is  much  larger  than  the  last,  approaching  nearly  in  its  dimen- 
sions the  Molgophis  macrurus.  It  is  represented  by  a  series  of  22  vertebrae,  which, 
like  those  of  Phlegethontia  linearis,  are  devoid  of  ribs,  abdominal  armature,  dilated 
neural  spines,  etc.  The  series  when  complete  must  have  been  very  long,  as  there  is 
little  difference  in  size  between  the  first  and  the  last  of  the  22.  They  are  marginate 
fore  and  aft,  and  much  contracted  medially,  owing  to  the  transverse  expanse  of  the 
diapophyses.  There  may  be,  indeed,  a  diapophysial  element  beneath  these,  but,  if 
so,  the  two  are  indistinguishable.  They  are  connected  by  longitudinal  impressions, 
indicating  the  existence  of  the  tendinous  bands  in  the  longitudinal  muscles  seen  in 
Amphiuma,  or  the  osseous  spicules  seen  in  the  same  situation  in  birds.  The  neural 
spines,  as  indicated  by  their  narrow  bases,  occupy  the  length  of  the  neural  arch,  and 
remind  one  of  Amphiuma.    Width  of  one  of  the  vertebrae,  3  lines. 


CHAPTER  XXI. 

THE  MICROSAURIAN  FAMILY  SAUROPLEURID^,  FROM  THE  COAL  MEASURES 

OF  OHIO. 

Family  SAUROPLETJRIDvE  Hay,  1902. 

Hay,  Bull.  U.  S.  Geol.  Surv.,  No.  179,  p.  419,  1902. 

The  present  family  is  an  association  of  related  forms  due  to  similar  structure 
of  vertebrae,  ribs,  ventral  scutellation,  and  limbs.  There  is  no  character  in  the  skull 
which  would  indicate  a  separation  of  the  genera  here  included,  at  least  in  the  light 
of  present  knowledge. 

The  family  may  be  characterized  as :  Subaquatic  or  terrestrial  vertebrates  with 
a  typically  amphibian  development  of  the  ventral  armature ;  ribs  intercentral,  as  in 
all  members  of  the  order;  skull  elongate  and  slender  or  broad  and  obtuse;  cranial 
and  dermal  elements  of  the  pectoral  girdle  sculptured;  lateral-line  canals  indicated 
in  one  genus,  Saurerpeton;  limbs  well  developed,  with  well-developed  digits  and 
ungual  phalanges  claw-like;  body  usually  slender,  broad  in  Saurerpeton;  the  ribs 
broad  and  heavy;  the  vertebrae  relatively  stout;  ventral  armature  highly  developed, 
reaching  the  height  of  specialization  among  the  Microsauria ;  scutellse  consisting  of 
rods,  plates,  or  stout  bristles.  The  family  is  represented  by  5  genera:  Sauropleura, 
Saurerpeton,  Ctenerpeton,  Leptophr  actus,  and  Eurythorax,  the  association  of  the  last 
two  genera  being  provisional.    The  genera  may  be  distinguished  as  follows: 

I.  Pectoral  elements  sculptured,  clavicles  triangular,  interclavicle  diamond-shaped,  ventral  scutellae  rods, 

arranged  en  chevron  with  anterior  angle Sauropleura 

II.  Pectoral  elements  slightly  sculptured,  cranium  broad,  obtuse  and  sculptured,  ventral  armature  broad 

imbricated  plates  extending  on  to  the  throat Saurerpeton 

III.  Limbs,  skull,  arches  and  dorsal  vertebra?  unknown,  caudal  vertebrae  with  fan-shaped  neural  and  haemal 

spines  which  may  indicate  relationship  with  Ptyonius  and  CEstocephalus,  but  in  those  genera  the 
ventral  armature  is  weakly  developed;  ventral  scutelte  curved  rod-like  plates  arranged  en  chevron 
with  anterior  angle,  marked  in  abdominal  region  by  distinct  rounded  pits Ctenerpeton 

IV.  Known  only  from  fragments  of  the  skull,  teeth  large  and  fluted;  association  in  family  provisional. .     Leplophractus 
V.  Known  only  from  a  single  interclavicle  of  peculiar  form  which  resembles  that  of  Saurerpeton;  association 

in  the  family  provisional Eurythorax 

The  members  of  this  family  are  confined  to  the  deposits  of  the  Coal  Measures 
at  Linton,  Ohio.  Ctenerpeton,  and  possibly  Sauropleura,  were  highly  developed 
swimmers,  but  the  strength  of  the  limbs  as  exhibited,  especially  by  Sauropleura  and 
Saurerpeton,  indicates  that  they  had  not  entirely  forsaken  the  land. 

Genus  SAUROPLEURA  Cope,   1868. 

Newberry,  Proc.  Phila.  Acad.  Nat.  Sci.,  vm,  p.  98,  1856  {Pygopterus  scutellalus). 
Cope,  Trans.  Am.  Phil.  Soc,  p.  22,  1869. 
Cope,  Geol.  .Surv.  Ohio,  it,  pt.  11,  p.  402,  1875. 
Hay,  Bull.  U.  S.  Geol.  Surv.,  No.  179,  p.  419,  1902. 

Type:  Sauropleura  scutellata  Newberry. 

Vertebrae  and  ribs  well  developed;  limbs  4,  rather  large;  5  digits  in  the  fore- 
foot; carpus  cartilaginous.  Ventral  armature  of  closely  arranged  rhomboidal 
scuta,  arranged  in  lines,  which  are  closely  placed  in  chevrons,  with  the  angle 
anterior. 

155 


I56  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH   AMERICA. 

In  none  of  the  species  of  this  genus  have  the  usual  3  thoracic  shields  (clavicles 
and  interclavicles)  been  observed.  The  abdominal  scuta?,  on  the  other  hand,  are 
much  like  those  of  Saurerpeton,  being,  however,  smaller. 

The  species  formerly  described  by  Cope  in  the  genus  Colosteus  are  included  in 
Sauropleura,  where  they  find  their  closest  allies. 

There  are  7  species  belonging  in  this  genus:  Sauropleura  digitata  Cope,  5.  new- 
berryi  Cope,  S.  foveata  Cope  {Colosteus),  S.  scutellata  Newberry  {Colosteus)  (type 
of  genus),  5.  pauciradiata  Cope  {Colosteus),  S.  longidentata  Moodie,  S.  enchodus 
Cope  {Anisodexis) .  The  species  described  by  Cope  as  Sauropleura  latithorax  is 
regarded  as  belonging  to  a  distinct  genus,  Saurerpeton. 

Sauropleura  scutellata  Newberry. 

Newberry,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  98,  1856  (Pygoplerus  sculellalus). 
Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  215,  1868. 
Cope,  Trans.  Am.  Phil.  Soc,  p.  22,  1869. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  402,  1875. 

Type:  Specimen  No.  8669  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.     (Plate  14,  fig.  3.) 

This  species  was  first  described  by  Newberry  as  a  fish  belonging  to  the  genus 
Pygopterus.  Cope  later  placed  it  under  the  genus  Colosteus  and  clearly  showed  its 
amphibian  characters.  The  genus  Colosteus  was,  however,  based  on  a  misconcep- 
tion. Cope  in  1897  and  Hay  (317)  referred  the  species  to  Sauropleura,  where  it 
is  retained.  The  species  is  represented  by  a  single  individual  preserved  on  a  block 
of  coal  from  Linton,  and  is  also  indicated  by  an  interclavicle  and  its  obverse;  this 
element  is  of  larger  size  than  that  of  the  type  and  was  referred  by  Cope  to  Colosteus 
pauciradiatus.  The  characters  of  the  plate  are,  however,  so  identical  with  those  of 
the  interclavicle  in  the  type  specimen  that  it  is  unhesitatingly  referred  to  the  pres- 
ent form. 

The  type  specimen  consists  of  the  supero-lateral  view  of  a  crushed  cranium  with 
the  anterior  part  of  the  body,  exhibiting  the  interclavicle  and  the  ventral  scutella- 
tion.  No  limbs  have  been  observed  in  this  species.  The  mandibles  are  crushed 
across  the  cranium  in  such  a  way  as  to  obscure  its  structure.  The  boundary  of  the 
left  orbit  is  doubtfully  determined  as  being  a  little  back  of  the  median  line  of  the 
skull.  There  are  small  teeth  present  on  the  mandibles,  but  their  number  can  not  be 
determined.  The  cranial  elements  are  sculptured  with  radiating  grooves  and 
ridges,  but  these  are  weakly  developed.  The  snout  is  broad  and  but  little  narrower 
than  the  base  of  the  skull.     (Plate  21,  fig.  5.) 

The  interclavicle,  somewhat  displaced,  is  the  only  element  of  the  pectoral 
girdle  preserved.  It  is  peculiar  in  the  possession  of  a  backward  extension  which 
shows  a  beveled  edge.  The  plate  is  ornamented  by  radiating  grooves  and  ridges 
which  are  strongly  developed.  The  larger  specimen  of  an  interclavicle  shows  the 
same  characters  as  the  one  described,  and  it  differs  only  in  being  about  twice  as 
large.    There  are  no  traces  of  limbs. 

The  ventral  armature  of  the  body  is  rather  weak  as  compared  to  that  of  Sau- 
ropleura pauciradiatus,  but  it  is  still  composed  of  closely  packed  scutes  arranged 
en  chevron.    The  character  of  the  ventral  armature  and  the  sculpturing  of  the  inter- 


HIE   MICROSAURIAN   FAMILY   SAUROPLEURIDyE.  157 

clavicle  are  taken  as  the  principal  specific  characters.  From  the  other  species  of 
the  genvis  the  present  form  differs  in  its  more  slender  elements  of  the  ventral  armor 
and  in  the  form  of  the  skull.    The  ribs  are  not  clearly  defined. 

Measurements. 

mm.  mm. 

Median  length  of  skull 70  Length  of  mandibular  tooth 1.5 

Posterior  width  of  skull 40  length  of  entire  specimen 150 

Anterior  width  of  skull 20  Width  across  belly,  maximum 47 

Length  of  jaw 50  Length  of  interclavicle 27 

Anterior  width  of  jaw 7  Width  of  interclavicle 13 

Posterior  width  of  jaw 12  Width  across  posterior  extension 4 

Sauropleura  digitata  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  216,  1868. 

Cope,  Trans.  Am.  Phil.  Soc.,  xiv,  p.  15,  1869. 

Cope,  Geol.  Surv.  Ohio,  n,  pt.  11,  p.  403,  pi.  xxxvii,  fig.  1,  1875. 

Type:  Specimen  No.  8004  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.     (Plate  20,  fig.  4.) 

This  species  is  represented  by  a  single  individual  which  is  quite  distinct  from 
other  members  of  the  genus.  The  specimen  has  been  spread  over  a  surface  of  the 
coal,  and  exhibits  ventral  armature,  dorsal  region  with  ribs,  and  anterior  and  pos- 
terior limbs.    Skull  and  caudal  region  not  present. 

The  ventral  armature  is  arranged  in  parallel  lines  directed  obliquely  forwards 
and  continuous  on  the  median  line,  forming  there  a  chevron.  The  individual  ele- 
ments are  oat-shaped,  and  acuminate  at  both  ends.  They  are  moderately  imbri- 
cate in  an  antero-posterior  section.  On  the  pectoral  region  between  the  fore  limbs 
the  series  of  scutellae  assume  different  directions,  forming  chevrons  directed  back- 
wards, and  forming  with  those  of  the  belly  a  complete  X. 

The  humerus,  ulna,  and  radius  are  rather  stout,  and  of  a  size  relative  to  the 
body,  as  in  common  types  of  existing  lizards;  the  ulna  and  radius  separate.  The 
carpus  is  cartilaginous;  the  digits  are  5  well-developed  fingers  having  phalanges  in 
the  following  numbers,  commencing  on  the  inside:  3,  4,  5,  6,  5.  The  last  phalanx  of 
the  second  is  obscured,  and  it  is  not  positive  that  the  number  is  as  given;  it  is  more 
probable  that  it  was  4  than  3.  The  outer  toe  was  more  slender  than  the  others; 
the  distal  phalanges  of  all  the  toes  are  stout,  as  in  modern  Caudata. 

The  ribs  are  long  and  curved  as  in  reptiles,  and  judging  by  their  distances  the 
vertebrae  are  short ;  the  latter  are  not  well-defined,  but  there  is  no  indication  of  prom- 
inent spines  of  any  kind.  The  pelvic  bones  and  portions  of  the  hind  limbs  are  pres- 
ent, but  so  obscured  and  confused  as  not  to  be  easily  made  out.  Enough  remains 
to  show  that  the  hind  limbs  were  longer  than  the  fore.  Thirteen  ribs  on  one  side 
and  twelve  on  the  other  are  preserved,  with  short  ribs  in  the  sacral  region.  The 
specimen  is  very  indistinct  and  it  is  difficult  for  one  to  be  sure  of  all  the  characters 
described  by  Cope. 

Measurements  of  the  Type  of  Sauropleura  digitata  Cope. 

mm.  mm. 

length  of  specimen 115            Length  of  radius 10 

Greatest  width 80            Length  of  ulna 9 

Diameter  of  ventral  scute .75      Length  of  metacarpal 4 

Length  of  rib 15    '        Length  of  fourth  digit  of  hand 22 

Width  of  rib I            Length  of  interclavicle 20 

Length  of  humerus 20           Width  of  interclavicle 1 1 


I58  THE   COAL   MEASURES    AMPHIBIA   OF   NORTH    AMERICA. 

The  interclavicle  is  diamond-shaped,  with  surface  punctate  and  edges  radiately 
grooved  at  a  distance  of  1.5  mm.  from  the  edge.  The  hand  on  the  right  side  of  the 
specimen  contains  3,  3,  3,  6,  3  phalanges.    The  vertebras  are  all  imperfectly  preserved. 

Other  specimens  of  this  species  are  2567  (48),  8376  G,  8704  G,  American  Museum 
of  Natural  History.  Coal  Measures  of  Linton,  Ohio.  Collected  by  Dr.  J.  S.  New- 
berry. 

Sauropleura  newberryi  Cope. 
Cope,  Geol.  Surv.  Ohio,  II,  pt.  II,  p.  404,  pi.  xxxvii,  figs.  2  and  3;  pi.  xli,  fig.  5,  1875. 

Type:  Specimen  No.  8612  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  type  specimen  of  the  species  exhibits  a  portion  of  the  posterior  part  of  the 
skull,  a  considerable  part  of  the  body,  with  the  fore  limbs  and  abdominal  scutellae. 
No  vertebrae  can  be  definitely  discovered  and  the  ribs  are  not  distinctly  visible. 
The  cranial  fragment  is  the  upper  surface  of  the  tabulare  and  adjacent  elements, 
and  a  broad  band  of  the  posterior  parts  of  these  is  seen  to  be  smooth,  and  is  pre- 
ceded by  a  slightly  roughened  surface.  The  abdominal  scutae  are  diamond-shaped, 
and  are  thin  and  light,  not  massive,  as  in  S.  scutellata,  and  are  sometimes  marked 
with  a  median  longitudinal  keel.  The  fore  limb  is  large,  especially  the  humerus, 
which  is  much  dilated  distally,  and  has  a  strong  crest  on  the  outer  side  from  near 
the  proximal  end.  The  ulna  and  radius  are  much  shorter,  and  more  dilated  prox- 
imally  than  distally;  they  are  well  separated.    No  phalanges  are  preserved.. 

The  species  is  represented  by  other  specimens,  all  of  which  are  unsatisfactory 
in  determining  the  structure  of  the  form.  A  skull,  apparently  complete,  is  crushed 
out  flat,  so  that  little  can  be  said  of  structure.  The  teeth  are  rather  long,  straight, 
acute,  and  striate  at  the  base.    The  orbits  are  long  and  narrowed  in  front. 

Measurements  of  Sauropleura  newberryi  Cope. 
(.No.  8612  G,  and  two  unnumbered  specimens,  American  Museum  of  Natural  History.) 

mm.  mm. 

Length  of  humerus 35  Width  at  anterior  angle  of  orbits 40 

Proximal  width  of  humerus 8  lnterorbital  space 13 

Distal  width  of  humerus 14  Width  of  orbit 13 

Length  of  ulna 19  Length  of  orbit 27 

Proximal  width  of  ulna 8  Length  of  a  maxillary  tooth 4 

Posterior  width  of  skull 67  Diameter  of  same  at  base 2 

Sauropleura  pauciradiata  Cope. 

Cope,  Trans.  Am.  Phil.  Soc.,  xv,  p.  275,  1874. 

Cope,  Geol.  Sun'.  Ohio,  n,  pt.  11,  p.  408,  pi.  xl,  figs.  1-2,  1875. 

Type:  Specimen  No.  8671  G,  and  obverse,  American  Museum  of  Natural 
History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  was  founded  on  a  median  and  a  lateral  plate  of  2  individuals.  The 
clavicle  is  here  associated  with  S.  scutellata  and  the  interclavicle  thus  remains  as 
the  type  specimen.  There  are  other  remains  associated  by  Cope  with  S.  scutellata 
which  are  here  shown  to  have  closer  affinities  with  the  S.  pauciradiata. 

The  clavicle  is  a  right-angled  triangle,  the  inner  and  thin  edge  concave  poste- 
riorly, the  posterior  convex.  The  ridges  and  grooves  are  well  developed  on  the 
specimen.    The  visceral  side  of  the  element  is  smooth. 


THE   MICROSAURIAN   FAMILY   SAUROPLEURID^E. 


159 


,<W%; 


Cope  referred  to  S.  scutellata  the  larger  part  of  an  individual  with  strongly 
developed  ventral  scutellation.  This  he  figured  on  plate  xxxvi,  fig.  2  (123). 
Even  a  cursory  glance  will,  however,  suffice  to  show  that  the  sculpturing  of  the 
clavicle  just  described  and  that  exhibited  by  the  specimen  there  figured  are  identical. 
Closer  examination  shows  important  differences  between  the  species,  the  principal 
distinctions  being  the  strongly  developed  ventral  scutellae  in  Sauropleura  scutellata, 
the  difference  in  form  and  sculpture  of  the  interclavicles,  and  the  posterior  extension 
of  the  interclavicle  in  5.  scutellata  which  is  wanting  in  S.  pauciradiata.  The  sculp- 
turing of  the  interclavicle  in  the  form  figured  by  Cope  as  S.  scutellata,  just  referred 
to,  is  identical  with  the  sculpture  of  the  clavicles,  as  would  be  expected.  The  ridges 
on  all  of  the  pectoral  elements  of  S.  pauciradiata  are  strong  and  are  rather  few  in 
number,  while  in  S.  scutellata  the  sculpturing  is  more  in  the  form  of  interrupted 
grooves. 

The  specimen  (plate  14,  fig.  3)  exhibits  the  great  part  of  the  body  with  one  fore 
limb.  The  skull  is  wanting.  The  belly  was  very  broad  and  strongly  protected  by 
broad,  long  scutes  arranged  en  chevron.  The  scutes  are  close  together  and  form  a 
compact  ventral  armor  for  the  animal.  The  fore  limb  is  very  weak.  The  humerus  is 
represented  by  its  distal  end  only.  The  radius  and  ulna  are  very  short  and  weakly 
developed  in  comparison  to  the  size  of  the  animal.  The  limbs  could  not  have  sup- 
ported the  animal  on  land  and  served,  probably, 
merely  as  organs  of  equilibration,  for  the  animal 
was  undoubtedly  aquatic.  The  fingers  are  not 
all  preserved  and  there  is  no  carpus. 

Numerous  other  remains  formerly  associated 
with  S.  scutellata  are  here  referred  to  S.  pauci- 
radiata, on  account  of  the  strongly  developed 
ventral  armor,  which  is  different  from  that  of 
the  type  of  5.  scutellata.  The  remains  do  not, 
however,  add  to  our  knowledge  of  the  anatomy 
of  the  forms,  as  they  are  very  fragmentary. 

Two  skulls  are  provisionally  associated  with 
this  species.  One  of  these  skulls  is  figured  by 
Cope  (123)  on  plate  xxxm,  fig.  1.  It  is  there 
referred  to  S.  scutellata.  That  it  can  not,  how- 
ever, be  referred  to  that  species  is  manifest  when 
the  teeth  are  observed.  In  the  type  of  S.  scutellata  the  teeth  are  very  small,  sharp 
denticles,  while  in  the  skull  under  discussion  the  teeth  are  well  developed  and  their 
bases  are  longitudinally  grooved.  The  teeth  are  elongate  in  the  anterior  part  of  the 
skull  and  are  shorter  posteriorly.  They  are,  however,  all  strong.  The  skull  is  acu- 
minate and  the  orbit  is  located  about  midway  of  its  length.  The  jaw  is  slightly  longer 
than  the  cranium.  The  structure  of  the  cranium  can  not  be  determined  in  either 
skull,  and  in  one  only  the  position  of  the  orbits  and  the  teeth. 


Fig.  34- 

A.  Interclavicle   of   Sauropleura   pauciradiata 

Cope.     X  1.    (After  Cope.) 

B.  Left  clavicle  of  Sauropleura  pauciradiata 

Cope.    X  1.    (After  Cope.) 


l6o  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

Measurements  of  the  Specimens  of  Sauropleura  pauciradiata  Cope. 

ram.  ram. 

Length  of  clavicle,  left 43      Space  between  ridges 2 

Width  of  clavicle 25 

Large  specimen  No.  8637  G  and  obverse  (13)  described  as  Colosteus  sculcllatus  Newb. 

mm.  ram. 

Length  of  specimen,  as  preserved 117  Width  of  clavicle 18 

Width  of  specimen,  maximum 61  Length  of  interclavicle 40 

Length  of  fore  arm 9  Width  of  interclavicle 23 

Length  of  hand 10  Width  of  a  ventral  scute 1.5 

Length  of  metacarpus 5  Length  of  scute  from  angle  to  end 35 

Length  of  clavicle 40 

Skull  No.  8666  C,  plate  xxiii,fig.  I  (Cope,  123),  Colosteus  scutellatus. 

mm.  mm. 

Length  of  skull,  as  preserved 65      Width  of  mandible,  maximum 10 

Width  of  skull 4°      Length  of  longest  tooth 5 

Diameter  of  orbit 6      Width  of  tooth  at  base 1.5 

Length  of  mandible 70 

Skull  Nus.  8602  G  and  860S  G. 

mm.  mm. 

Length  of  skull,  as  preserved 50      Diameter  of  orbit 15 

Width  of  skull 60      Length  of  mandibular  tooth 4.5 

Other  specimens  associated  with  this  species:  Nos.  8668  G,  86740,85540, 
8661  G.    All  specimens  in  the  American  Museum  and  all  from  Linton,  Ohio. 

SCUTES    OF    SAUROPLEURA. 

There  are  found  associated  with  the  remains  of  the  genus  Sa  u ro pleura  a  num- 
ber of  heavy  scutes  or  scales.  There  are  three  of  them  on  the  same  block  as  the 
specimen  of  S.  longidentata  and  are  provisionally  referred  to  the  genus.  There 
are  a  number  of  scutes  preserved  separately,  but  they  agree  in  their  characters 
with  those  discovered  on  the  specimens.  The  scutes  are  elongate  and  usually  acu- 
minate at  one  end  and  having  a  broad  base  at  the  other.  The  acuminate  end  is 
slightly  bent  to  one  side  so  as  to  present  the  appearance  of  a  hook.  Others  are 
shield-shaped  and  are  quite  large,  while  the  majority  of  the  hook-shaped  ones  are 
small.  The  shield-shaped  elements  have  a  rounded  boss  near  the  center  of  the 
plate  and  the  edges  are  imbricated.  Their  nature  and  their  proper  location  on  the 
animal  are  a  puzzle.  They  may  not  belong  to  the  genus,  but  have  been  noticed  with 
the  remains  of  at  least  3  species. 

Measurements  (in  Millimeters)  of  Scutes  Associated  with  the  Specimens  of  Sauropleura. 

No.  4513  U.  S.  National  Museum: 

Length 27 

Width,  maximum 14 

Width,  minimum 3 

Scutes  associated  with  the  specimen  of  Sauropleura  longidentata: 

Width  across  base 3 

Width  across  tip 1 

Additional  specimens  are:  Nos.  3,  8673  G  and  8470  G,  American  Museum  of 
Natural  History. 

Sauropleura  longidentata  Moodie. 
Moodie,  Jour.  Geol.,  17,  No.  1,  pp.  74-76,  figs.  18,  19,  1909. 

Type:  Specimen  No.  8619  G,  American  Museum  of  Natural  History. 
Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  species  may  be  distinguished  from  other  members  of  the  genus  by  the  large 
size  and  shape  of  the  cranium  (462)  and  the  broad  mandible  (plate  16,  figs.  2,  3) 


MOODII 


FUkTS  22 


1. 


I. 


Type  of  Leplophractns  lineolatus  Cope,  from  the  Coal  Measures  of  Linton,  Ohio.  Por- 
tions of  maxilla  and  mandible  of  left  side  with  teeth.  XI.  Original  in  American 
Museum  of  Natural  History. 

Type  of  Proterpelon  gurleyi  Moodie,  from  the  Coal  Measures  of  Illinois,  near  Danville. 
Cervical  of  an  otherwise  unknown  vertebrate.  Neural  spine  to  the  right. 
Original  in  Walker  Museum,    University  of  Chicago     X  2. 

Amphibian  phalanx  from  the  Coal  Measures  near  Breeze,  Illinois,  of  an  unknown 
species.     The  probable  form  of  the  element  is  represented.     X  3. 

Large  rib  of  a  stereospondylous  stegocephalan  otherwise  unknown.  The  rib  may 
represent  a  species  of  Macrerpeton  or  may  even  belong  with  Macrerpeton  deani 
Moodie,  but  exact  identification  will  have  to  wait  for  future  discoveries.  From 
the  Coal  Measures  of  Linton,  Ohio.     Original  in  U.  S.  National  Museum.     X  1. 

Type  of  Cope's  species  Tia/i/aniis  mordax  referred  by  him  to  the  cranium,  on  account 
of  the  sculpturing  of  the  elements.  We  now  know  the  specimen  to  be  portions  of 
the  interclavicle  and  clavicles  of  Diceralosaiirits  ptinctolineattis,  as  Cope  suggested 
they  might  be.     Original  in  American  Museum  of  Natural  History.     X  1. 

Skull  of  Raphetes  planiceps  Owen  from  the  Coal  Measures  of  Nova  Scotia.  0.45. 
Original  in  the  British  Museum,  South  Kensington.     After  Owen. 


THE   MICROSAURIAN   FAMILY   SAUROPLEURIM:.  l6l 

with  its  very  long  teeth.  The  skull  of  Sauropleura  digitata  Cope  is  not  known,  but 
the  body  of  that  animal  as  preserved  represents  far  too  small  a  form  for  the  skull  to 
be  referred  to  that  species.  The  skull  is  fully  half  as  long  as  the  dorsal  region  of  S. 
digitata  Cope,  so  that  an  association  of  the  remains  would  be  incongruous.  It  differs 
from  the  skull  of  S.  scutellata  Newberry  in  size  and  proportions.  The  skull  of  S. 
scutellata  is  narrow,  while  in  S.  longidentata  it  is  quite  broad.  The  teeth  of  the  latter 
are  also  characteristic  of  the  species,  since  in  all  other  known  species  of  this  genus  in 
which  the  skull  is  preserved  the  large  anterior  tooth  is  wanting. 

The  bones  of  the  skull  show  the  coarse  sculpturing  of  the  larger  species  of  Micro- 
sauria.  It  consists  more  of  radiating  grooves  than  of  pits.  The  skull,  as  restored 
(462),  is  broadly  ovate,  with  the  posterior  border  truncate.  The  muzzle  is  broad 
and  the  nostrils  are,  apparently,  located  near  the  anterior  margin. 

The  posterior  border  of  the  orbits  lies  near  the  median  transverse  line  of  the 
skull.  They  are  circular  and  are  removed  some  distance  from  the  margin  of  the 
cranium.    Only  the  frontal  and  parietal  can  be  determined  with  certainty. 

The  mandible  is  heavy  and  is  provided  with  pleurodont,  heterodont  teeth.  Near 
the  anterior  end  of  the  mandible  there  is  a  very  long  fang-like  tooth,  longitudinally 
striated  and  slightly  recurved,  which  arises  from  a  broad  base  and  attains  to  con- 
siderable prominence.  The  other  teeth  are  smaller,  though  the  next  succeeding  one 
is  still  of  considerable  size.  All  of  the  teeth  preserved  are  longitudinally  striated, 
but  only  the  two  anterior  ones  are  recurved  to  any  extent. 

Measurements  of  the  Type. 

mm.  mm. 

Length  of  the  skull  in  median  line 75  Width  of  jaw,  maximum 16 

Width  of  skull  at  posterior  border,  estimated 80  Width  of  jaw,  minimum 5 

Width  of  skull  across  orbits 60  Length  of  largest  tooth 11 

Width  of  orbit 10.6  Width  of  longest  tooth  at  base 4.5 

Length  of  orbit 12  Length  of  shortest  tooth 3 

Interorbital  space 16  Width  of  shortest  tooth  at  base 1 

Length  of  jaw,  as  preserved 48 

Sauropleura  foveata  Cope. 

Cope,  Trans.  Am.  Phil.  Soc.,  xiv,  p.  24,  1869. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  406,  pi.  xxxvi,  fig.  1,  1875. 

Type :  Specimen  No.  8676  G  and  obverse  No.  8675  G,  American  Museum  of 
Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  species  is  represented  by  an  interclavicle  and  its  impression,  which  show  a 
beautiful  sculpturing  entirely  distinct  from  that  of  the  other  species  of  this  genus. 
In  size  it  is  intermediate  between  the  largest  of  the  interclavicles  of  S.  scutellata  and 
S.  pauciradiata.  The  pattern  of  the  sculpturing  is,  however,  its  main  distinction. 
The  plate  is  finely  pitted  and  there  are  few  evidences  of  grooves.  Near  the  posterior 
border  of  the  plate  the  pits  become  somewhat  defined  by  ridges  which  take  on  a 
radiating  pattern  with  the  center  of  the  plate  as  the  center.  The  beveled  margins 
are  rugose,  except  at  the  edges. 

Measurements  of  the  Type  Specimen  of  Sauropleura  foveata  Cope. 

mm. 

Median  length  of  interclavicle 43 

Width  of  interclavicle,  maximum 23 


1 62  THE   COAL    MEASURES   AMPHIBIA    OF    NORTH    AMERICA. 

Sauropleura  enchodus  Cope. 
Cope,  Proc.  Am.  Phil.  Soc,  p.  406,  1885  (Anisodexis). 

Type:  Specimen  No.  (51)  2558,  American  Museum  of  Natural  History,  Newberry 
Collection. 

Horizon  and  locality:  Discovered  by  Sam  Huston  at  the  Linton,  Ohio,  Coal 
Measures. 

An  examination  of  the  type  specimen  (plate  16,  fig.  4)  of  this  form  resulted  in  no 
new  facts.  The  reference  of  the  species  to  the  Permian  genus  Anisodexis  by  Cope 
is  probably  incorrect.  Our  knowledge  of  the  two  known  species  of  this  genus  is  not 
sufficient  to  separate  them,  but  for  the  sake  of  convenience  the  Linton  species  is 
placed  in  the  genus  Sauropleura.  It  is  located  here  principally  on  account  of  the 
form  and  structure  of  the  teeth.     Cope's  original  description  is  given  below: 

"The  generic  characters  are  apparent  in  the  very  unequally  sized  teeth  with  round 
section.  The  portion  upon  which  the  species  is  based  is  a  part  of  the  right  ramus  of  the 
mandible,  which  is  in  the  specimen  viewed  from  the  inner  side.  The  jaw  is  obliquely  and 
smoothly  truncated  from  below,  for  the  symphysis  and  surface  of  the  bone  is  smooth. 
There  is  a  very  large  tooth  near  the  extremity  of  the  dentary  bone.  Behind  it  is  an  interval 
equal  to  three  times  the  diameter  of  its  base,  which  is  followed  by  a  tooth  of  about  one- 
third  the  length  of  the  first  tooth.  Posterior  to  this  one  are  two  teeth  of  the  same  size  as 
the  second,  all  being  separated  from  each  other  by  about  a  tooth's  diameter.  These  are 
followed  by  three  subequal  teeth  of  about  two-thirds  the  length  of  the  first  tooth,  and 
separated  by  about  their  own  diameter  from  each  other.  They  are  all  perfectly  straight, 
very  acute,  and  without  the  trace  of  a  cutting-edge.  The  inflection-grooves  extend  to  or 
a  little  beyond  the  middle  of  the  length." 

The  present  species  is  smaller  than  the  type  of  Anisodexis  imbrecarius  from  the 
Texas  Permian,  to  which  genus  Cope  originally  referred  the  present  species,  and  the 
apices  of  the  teeth  do  not  display  the  opposite  cutting-edges  seen  in  the  Texas  form. 

Measurements  of  the  Type  Specimen. 

mm.  mm. 

Length  of  jaw,  including  7  teeth 31  Length  of  third  tooth 3.5 

Depth  of  ramus  at  second  tooth 10         Length  of  sixth  tooth 7.5 

Length  of  first  tooth 10.5 

SKIN    OF    SAUROPLEURA    SP. 
Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  p.  355,  pi.  lx,  fig.  1,  1909. 

The  present  specimen  is  interesting  on  account  of  the  presence  of  what  I  take  to 
be  a  portion  of  the  skin,  which  is  preserved  as  a  smooth  mold  over  the  ribs  and  ven- 
tral scutellas.  The  skin  was  undoubtedly  that  of  the  back,  since  the  creature  is 
preserved  on  its  belly,  and  is  interesting  in  not  showing  the  slightest  trace  of  scales 
or  other  hard  plates.  The  ventral  scutellae  are  characteristic  of  the  species  of  the 
genus  Sauropleura.  With  one  species  of  this  genus,  Sauropleura  scutellata  Newberry, 
the  writer  has  found  associated  scutes  of  some  size,  and  the  same  fact  has  been 
noted  by  Cope. 

Genus  SAURERPETON  Moodie,  1909. 
Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  80,  fig.  23,  1909. 

Type:  Saurerpeton  latithorax  Cope. 

This  generic  name  is  erected  for  the  reception  of  a  single  species  described  by 
Cope  in  1897.  The  name  is  made  necessary  by  the  wide  divergence  of  the  charac- 
ters exhibited  by  the  present  species  from  those  of  the  species  of  the  genus  {Sauro- 


THE   MICROSAURIAN   FAMILY   SAUROPLEURID^E.  163 

pleura)  to  which  Cope  (176)  referred  this  species.    The  form  is  not  a  member  of  the 
genus  Sauropleura,  for  reasons  given  below. 

The  species  of  the  genus  Sauropleura  have  a  lanceolate  head  with  homodont  den- 
tition or  nearly  so.  The  orbits  are  located  well  back  in  the  skull.  The  form  of  the 
body  is  elongate  and  slender  and  the  limbs  where  known  are  long  and  attenuated. 
The  ventral  scutellation  consists  of  oat-shaped  scutes  arranged  in  a  chevron  series. 
The  form  here  described  as  Saurerpeton  latithorax  Cope  has  nearly  the  opposite  of 
all  of  these  characters,  and  it  is  incongruous  to  locate  the  form  under  the  former 
genus.  The  skull  of  Saurerpeton  latithorax  Cope  is  broad  and  heavy.  The  teeth 
are  heterodont.  The  body  is  broad  and  stout  and  the  limbs  are  of  unusually 
strong  proportions.  The  character  of  the  ventral  armature  is  also  of  a  very  differ- 
ent type.  In  Saurerpeton  it  consists  of  very  broad  imbricating  scutes  which  form  a 
single  piece  across  the  abdomen  and  are  angulated  to  form  the  chevron  pattern 
which  is  so  common  among  the  Stegocephalia. 

Saurerpeton  latithorax  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  xxxvi,  p.  86,  pi.  iii,  fig.  4,  1897. 
Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  80,  fig.  23,  1909. 

Type:  Specimen  No.  4471,  U.  S.  National  Museum. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.    Collected  by  R.  D.  Lacoe. 

This  species  is  indicated  by  remains  of  the  anterior  half  of  a  large  amphibian 
(plate  17)  preserved  on  a  block  of  bituminous  coal  from  the  Linton  mine.  The  form 
is  unusual  in  the  proportions  of  the  head  and  the  width  of  the  thoracic  region.  In 
these  characters  it  stands  alone  among  the  Amphibia  from  this  locality,  where  the 
forms  are  for  the  most  part  of  rather  slender  build  and  tapering,  pointed  head. 

The  skull  is  represented  in  a  fairly  complete  condition  and  shows  the  usual 
stegocephalian  arrangement  of  the  skull  elements,  as  well  as  the  sculpturing  of  the 
bones,  which  is  similar  to  that  found  in  other  members  of  the  Microsauria.  The 
skull  is  broadly  rounded,  with  the  posterior  border  incised,  the  broad  tympanic 
notches  thus  rendering  the  shape  of  the  skull  somewhat  like  that  of  the  branchio- 
saurs.  The  orbits  are  broad  ovals  and  lie  well  forward  in  the  skull.  They  are  sepa- 
rated by  a  space  which  is  greater  than  the  greatest  diameter  of  the  orbit.  The 
pineal  foramen  lies  well  back  and  is  clearly  indicated  as  a  circular  opening  which 
lies  in  the  median  suture  in  the  posterior  half  of  the  parietals.  The  nostrils  seem  to 
be  elongate  and  have  an  oblique  position,  as  is  represented  in  the  diagram  (fig.  35), 
but  this  character  is  not  ascertained  definitely. 

The  borders  of  the  premaxillae  and  the  anterior  suture  of  the  nasal  can  not  be 
determined,  though  they  may  have  had  some  such  arrangement  as  suggested.  The 
nasal  is  represented,  so  far  as  is  determinable,  by  an  oblong  element  lying  between  the 
frontal  and  the  anterior  border  of  the  skull.  The  frontals  are  very  large  and  form  a 
portion  of  the  inner  border  of  the  orbit.  The  parietal  is  probably  the  largest  bone 
in  the  cranium  and  together  the  two  elements  form  a  large  quadrangular  space 
in  the  posterior  half  of  the  skull.  They  inclose  the  circular  pineal  foramen.  The 
postparietal  is  a  small  element  lying  on  the  posterior  border  of  the  skull  and  with 
the  tabulare  and  a  part  of  the  supratemporal  forms  the  projection.    The  prefrontal 


1 64 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


is  probably  a  small  element,  especially  if  the  lacrimal  is  present.  There  seems  to 
be  an  indication  of  a  suture  separating  the  lacrimal  from  the  prefrontal,  but  this 
character  is  not  assured.  The  boundaries  of  the  maxillae  are  clear  anteriorly.  They 
indicate  that  this  element  was  elongate,  as  is  usual,  and  impressions  of  teeth  borne 
on  the  mandible  would  indicate  the  probability  that  the  teeth  were  heterodont. 
In  the  premaxillary  region  there  is  a  long,  strong  tooth  preserved,  and  on  the  max- 
illary near  the  posterior  ex- 
tremity of  this  bone  there  are 
impressions  of  teeth  which  are 
no  more  than  one-fourth  as 
large  as  the  premaxillary  one. 
The  borders  of  the  jugal  can 
not  be  ascertained,  since  the 
skull  is  injured  on  both  sides 
in  this  region.  Likewise  the 
quadratojugal  is  conjectural. 
The  boundaries  of  the  post- 
frontal  and  the  postorbital  are 
clearly  ascertained.  They  to- 
gether form  the  entire  poste- 
rior border  of  the  orbit  and 
send  prolongations  along  the 
lateral  borders  of  these  open- 
ings. They  are  both  acumi- 
nate posteriorly  and  these 
points  are  inclosed  by  the  tab- 
ulare  and  supratemporal  for 
the  postfrontal  and  by  the 
squamosal  and  prosquamosal  „ 

f       ,,                      ,  .      .       _,.  riG.  35. — Outline  drawing  of  the  skull  and  skeleton,  as  preserved,  of 

Space  IOr  tne  pOStOrbltal.      I  he  Saurerpelon  latithnrax  (Cope),  from  the  Linton,  Ohio,  Coal  Measures, 

hrainrlormo      ^f     +U^      4-„u    1  Original  in  U.  S.  National  Museum.     X  0.7s. 

boundaries     Ot     the      tabulare  Skull:  /,,  frontal;  j   jugal;   mx,  maxilla;  n,  nasal;  no,  nostril;  or,  orbit; 

Show   this  element  to  be  Quite  pnr'c  Paneta':  P°f,  postfrontal;  po,  postorbital;  pp,  postparietal;  pf, 

"  prefrontal;  pmx,  premaxilla;  qj,  quadratojugal;  m,  mandible;  spo, 

large    and    extending    forward  supraorbital   lateral-line  canal;  spt,  supratemporal;  sq,  squamosal; 

•     ,                             ...             ....  tub,  tabulare;  v,  pineal  foramen. 

mtO  an   aCUmmatlOn  Which    IS  Skeleton:  d,  clavicle;    c,  carpus;    rb,  ribs;  ft,  humerus;  it,  interclavicle; 

inclosed    bv   the    Oarietal    nnrl  »ie^metacarpals;/w,  pectoral  scutelte;r,  radius;  w,  ventral  scutclUe; 

the  postfrontal.  The  sutures  bounding  the  squamosal  have  been  obscured  by  injury 
in  removing  the  specimen  and  are  indeterminable. 

The  vertebral  column  is  represented  by  a  ridge  showing  along  the  median  plane 
of  the  specimen.  Its  characters  can  not  be  ascertained.  There  are  a  few  evidences 
of  ribs.  They  represent  long,  slender,  non-alate  elements,  their  mode  of  articu- 
lation with  the  vertebras  being  obscured  by  the  plate-like  ventral  armature  which 
covered  the  abdomen  of  the  animal. 

The  pectoral  girdle  is  represented  by  the  remains  of  three  elements  which  are 
interpreted  as  being  the  interclavicle  and  the  two  clavicles.  The  interclavicle  is 
broad  and  is  rounded  posteriorly.    There  is  no  evidence  of  the  usual  acumination 


THE   MICROSAURIAN   FAMILY   SAUROPLEURID/E.  1 65 

posteriorly.  The  element  is  nearly  as  wide  as  long.  There  is  a  prominent  longi- 
tudinal keel  on  the  ventral  surface  of  the  interclavicle  and  radiating  lines  which 
may  indicate  the  courses  of  blood-vessels  or  nerves  or  may  be  the  ornamentations 
of  the  element,  probably  the  latter.  The  clavicle  has  the  usual  microsaurian  form. 
It  has  three  points  and  is  truncate  exteriorly.  It  is  ornamented  with  radiating 
grooves  of  a  shallow  and  not  strongly  pronounced  character.  There  is  no  evidence 
of  the  coarse  sculpture  of  the  later  forms.  If  the  scapula  is  represented  it  is  merely 
by  an  indeterminate  fragment  insufficient  for  description. 

The  pectoral  limbs  are  preserved  nearly  entire.  The  left  fore  limb  lacks  only  a 
few  phalangeal  bones,  and  these  were  preserved  with  the  remainder  of  the  skeleton 
but  were  lost  in  the  mining  process.  The  humerus  is  an  extraordinary  element  on 
account  of  its  robust  dimensions.  It  is  very  stoutly  built  and  represents  a  power- 
ful limb.  It  is  expanded  at  each  extremity  and  the  width  of  its  shaft  is  about  equal 
to  one-fourth  of  its  length.  The  ulna  and  radius  present  the  same  characters  as 
the  humerus,  i.e.,  in  being  robust,  with  stout  shaft  and  expanded  ends.  The  ulna  is 
slightly  longer  than  the  radius  and  has  an  expanded  upper  end.  The  radius  is  short 
and  does  not  have  the  proximal  expansion.  The  carpus  was  cartilaginous.  Its  posi- 
tion is  represented  by  a  blank  space  on  the  coal.  There  are  4  digits  preserved  and 
in  all  probability  this  was  the  entire  number.  The  metacarpals  are  elongate  and 
expanded  at  the  extremities.  The  first  and  second  digits  are  represented  nearly  com- 
plete. The  first  digit  is  extremely  interesting  in  the  possession  of  a  claw-like  terminal 
phalanx  which  much  resembles  that  of  some  lizards.  There  are  3  phalanges  in  the 
first  digit  and  4  in  the  second.    The  phalangeal  formula  may  have  been  3-4-4?. 

The  ventral  scutellation  of  this  species  is  of  an  unusual  character.  It  consists 
of  broad,  imbricated  scutes  which  are  in  a  single  piece  and  which  are  arranged  in 
the  usual  chevron  pattern.  The  scutes  were,  apparently,  broadest  in  the  middle 
and  tapered  somewhat  at  the  extremities.  This  character  alone  is  sufficient  for 
separating  the  genus  from  that  of  any  other  known  form. 

The  genus  finds  its  nearest  allies  in  the  forms  of  the  species  of  the  genus  Sauro- 
plcitra,  in  which  Cope  formerly  located  the  present  species.  The  skull  of  the  form 
described  as  Tuditanus  radial  us  Cope  is  quite  similar  to  the  present  form,  both  in 
the  sculpturing  and  arrangement  of  the  elements.  The  characters  wherein  the  pres- 
ent form  resembles  the  species  of  Sauropleura  are  the  possession  of  broad  pectoral 
plates  and  strong,  digitate  limbs.  The  general  form  of  the  body  and  skull  is  different 
in  the  two  groups.  It  is  slender  in  Sauropleura  and  decidedly  stout,  short,  and 
heavy  in  Saurerpeton. 

M  KASI  KEMENTS  OF   THE    TYPE. 

mm.  mm. 

Length  of  specimen 130      Width  of  clavicle 18 

Median  length  of  skull 51       Length  of  abdominal  scutes 28 

Width  of  skull  at  posterior  border 62      Length  of  humerus 19 

Width  of  skull  across  orbits 48      Width  of  humerus  at  upper  end 6.5 

Length  of  orbit 14      Length  of  ulna II 

Width  of  orbit 10      Length  of  radius 9 

Interorbital  space 16      Length  of  metacarpal 3 

Length  of  longest  tooth  preserved 4       Length  of  first  digit 12 

Length  of  shortest  tooth  preserved I       Length  of  terminal  phalanx 3.5 

Length  of  the  interclavicle 26      Length  of  lower  jaw  on  the  curve 70 

Width  of  interclavicle,  maximum 23       Width  of  lower  jaw,  maximum 8 

Length  of  clavicle 22       Length  of  rib 30 


1 66  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

Genus  CTENERPETON  Cope,  18Q7. 
Cope,  Proc.  Am.  Phil.  Soc,  xxxvi,  p.  83,  pi.  iii,  fig.  1,  1897. 

Type :  Ctenerpeton  alveolatum  Cope. 

The  genus  Ctenerpeton  was  founded  by  Cope  for  the  reception  of  a  single  species 
which  presents  some  very  unusual  characters.  The  form  shows  close  relationships 
to  the  genera  Urocordylus  (334),  CEstocephalus,  and  Ptyonius.  These  genera  agree 
with  the  present  one  in  the  possession  of  very  characteristic  vertebrae  which  are 
signalized  by  the  elongate  and  ornamented  characters  of  the  neural  and  haemal 
spines.  These  project  prominently  from  the  body  of  the  vertebra  and  have  the 
ends  of  the  projections  truncate  and  divided  into  fine  points,  thus  causing  the  spine 
to  have  much  the  appearance  of  a  comb.  The  surface  of  the  neural  spine  is  some- 
times marked  with  a  shallow  groove.  The  spines  are  longer  and  more  slender  in 
the  genera  CEstocephalus,  Urocordylus,  and  Ctenerpeton  than  they  are  in  the  species 
of  the  genus  Ptyonius,  where  they  are  short,  although  the  usual  pectinations  are 
present. 

The  character  on  which  this  genus  rests  is  the  shelf -like  extension  (plate  23,  fig.  2) 
of  the  abdominal  plates.  This  is  of  a  very  unusual  character  and  entirely  unknown 
in  any  other  species  of  Carboniferous  Amphibia.  The  term  Ctenerpeton  has  reference 
to  the  fact  that  the  ends  of  these  shelf-like  plates  project  in  a  pectination  along  the 
side  of  the  abdomen. 

Ctenerpeton  alveolatum  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  xxxvi,  p.  83,  pi.  iii,  fig.  1,  1897. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  37,  p.  24,  pi.  10,  1909. 

Type:  Specimen  No.  4475,  U.  S.  National  Museum,  Lacoe  Collection. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  rests  on  a  single  specimen  (plate  19)  from  Linton,  Ohio,  and  is  pre- 
served on  a  block  of  bituminous  coal.  It  is  in  a  very  good  state  of  preservation. 
There  are  present  on  the  block  of  coal  a  part  of  the  right  fore  limb,  the  greater  part 
of  the  dorsal  portion  of  the  animal,  and  the  anterior  part  of  the  tail.  There  are  no 
evidences  of  hind  limbs,  although  this  may  not  be  taken  as  evidence  that  they  were 
not  present  on  the  animal.  No  thoracic  plates  have  been  observed.  Thexhevron 
armature  is  present  beyond  the  cloacal  region,  but  there  are  no  evidences  of  the 
specialized  clasping  organs  which  are  apparently  developed  from  the  abdominal 
armature  in  some  forms  (251). 

Each  dermosseous  rod  of  the  abdominal  scutellation  consists  of  three  pieces — 
a  median  angulated  portion  and  the  two  lateral  parts  which  form  the  shelf-like 
projection  along  the  side  of  the  abdomen  (plate  23,  fig.  2).  The  marginal  chevron 
differs  in  form  from  the  other  plate,  aside  from  the  fact  that  it  is  not  angulated. 
The  lateral  shelf  is  composed  of  flattened  plates  which  articulate  with  the  median 
piece,  and  at  the  place  of  articulation  there  is  a  ridge  present  in  the  specimen.  The  ex- 
terior plates  are  curved  backwards  and  are  somewhat  attenuated  distally.  They  are 
broader  than  the  median  piece  and  differ  also  in  the  absence  of  the  characteristic 
alveoli.  The  median  plate  is  angulated  and  is  of  more  slender  proportions  than  the  lat- 
eral pieces.     Its  ventral  surface  is  ornamented  with  a  single  row  of  closely  placed 


MOOOli 


PiATE  23 


2. 


Left  leg  and  pelvis  of  Ichlhynviihus  platypus  Cope,  from    the   Coal    Measures   of    Ohio. 

X  1.7.     Original  the  property  of  the  Department  of  Geology  of  Columbia  University. 

r=centrale;  /c"=fibula;yf=fibulare:  /v— -femur;  t=intermedium;  //=ilium;  7~=tibia; 

/,  /-5=distal  tarsalia;  /;'=tibiale:  F=caudal  vertebra;  /-K=digits. 
Ventral  scutellas  of  Ctenerpeton  alveolatum  Cope,  from  the  Coal  Measures  of  Ohio.     X  3.5. 

Original  in  U.  S.  National  Museum. 


THE   MICROSAURIAN   FAMILY   SAUROPLEURID.E.  1 67 

alveoli  which  resemble  in  a  great  degree  the  alveoli  of  the  jaw  of  some  small  animal. 
The  ventral  scutellation  is  broad  anteriorly,  but  becomes  more  slender  posteriorly 
and  shortly  posterior  to  the  cloacal  region  disappears. 

The  fore  limb  is  represented  by  the  upper  portions  of  the  ulna  and  radius  and 
2  digits  of  the  right  hand.  The  digits  are  long  and  slender  and  seem  to  represent 
digits  I  and  II,  since  they  show  evidences  of  3  and  4  phalanges  respectively.  The 
portions  of  the  fore  arm  preserved  are  too  meager  for  description. 

The  vertebrae  have  already  been  characterized  as  of  the  type  first  described  in 
Urocordylus.  The  neural  fans  are  not  much,  if  any,  wider  than  the  haemal  fans. 
They  are  both  situated  on  an  elongate  spine  with  a  slender  base.  The  edges  of  the 
two  fans  are  pectinated  and  the  dorsal  spine  is  distinguished  by  the  presence  of  a 
longitudinal  groove  in  the  center  of  the  spine.  The  length  of  the  tail  may  have 
been  considerable,  judging  from  the  character  of  the  vertebrae  preserved 

Measurements. 

mm.  mm. 

Length  of  specimen 150  Width  of  the  same  vertebra  with  spines 20 

Width  of  belly,  maximum 28  Height  of  neural  spine 8 

Length  of  lateral  chevron  plate 7  Height  of  haemal  spine 8 

Width  of  lateral  chevron  plate 1.75  Width  at  distal  end  of  hsemal  spine 3 

Length  from  tip  of  lateral  chevron  to  median  Width  at  distal  end  of  neural  spine 3.5 

angle 10  Length  of  foot,  as  preserved 12 

Length  of  a  caudal  vertebra 10 

Genus  LEPTOPHRACTUS  Cope,  1873. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  340,  1873. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  399,  1875. 
Cope,  Proc.  Am.  Phil.  Soc,  xx,  p.  461,  1882. 

Type:  Leptophractus  obsoletus  Cope. 

The  genus  was  established  on  various  parts  of  the  cranium  of  a  large  amphibian. 
The  only  parts  which  can  with  certainty  be  referred  to  the  genus  are  the  upper  and 
lower  jaws  of  3  specimens.  These  bear  large  teeth,  round  in  section  at  the  base, 
but  with  acute  compressed  apex,  with  a  cutting -edge  on  the  anterior  face;  the 
enamel  is  delicately  grooved,  as  an  external  indication  of  the  labyrinthic  structure. 
A  characteristic  feature  is  seen  in  the  presence  of  a  large  elongate  tooth  in  the  upper 
jaw,  in  the  position  of  a  canine  which  much  exceeds  in  length  any  of  the  others. 
The  sculpture  of  the  cranium  is  but  little  marked  in  the  known  specimens.  In  the 
type  the  lower  jaw  is  marked  with  inosculating  grooves.  Three  species  are  known, 
which  are  among  the  largest  of  the  Linton  Amphibia. 

Leptophractus  obsoletus  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  340,  1873. 
Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  399,  1875. 
Cope,  Proc.  Am.  Phil.  Soc,  xx,  p.  461,  1882. 

Type:  Specimen  Nos.  55  G  and  57  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  species  was  about  as  large  as  an  adult  Florida  alligator,  and  probably 
exceeded  or  at  least  equaled  in  size  any  of  the  Carboniferous  Amphibia.  The  fol- 
lowing account  is  taken  directly  from  Professor  Cope's  "  Batrachia  of  the  Ohio  Coal 
Measures"  (123).  The  description  has  been  verified  from  an  examination  of  the 
type  material. 


j68  the  coal  measures  AMPHIBIA  OF  NORTH  AMERICA. 

"  The  teeth  are  rather  distantly  grooved  for  some  distance  above  the  base.  They  are 
of  different  sizes;  the  smaller  are  compressed,  and  with  fore  and  aft  cutting  edges.  The 
external  surface  of  the  dentary  bone  is  marked  with  short  oblique  grooves  along  its  middle 
region;  above  these  are  grooves  which  inosculate,  forming  a  figure  like  an  open  net  dragged 
in  the  long  direction. 

"Excepting  the  grooves  the  teeth  are  smooth.  The  smaller  ones  are  close  together 
and  their  crowns  are  curved  backwards;  the  larger  ones  are  at  more  remote  intervals;  both 
have  enlarged  bases.  Whether  both  forms  are  in  the  same  series  I  can  not  determine. 
There  are  from  four  to  five  of  the  smaller  to  an  inch. 

Measurements  of  Type  of  Leptophractus  obsoletus  Cope. 

mm.  mm. 

Deplh  of  fragment  of  jaw 75      Width  of  paired  median  scuta 56 

Length  of  smaller  teeth 19      Width  of  single  scute 36 

Length  of  longer  tooth 23      Length  of  single  scute 48 

Width  of  vertex  at  middle  scuta 176 

' '  Some  vertebrae  were  found  at  the  same  locality,  but  there  is  no  evidence  as  to  the 
species  to  which  they  may  have  pertained.  They  are  short,  concave  on  one  end,  and 
probably  so  on  the  other.  The  centrum  of  one  is  12  mm.  in  diameter ;  neural  spines  injured." 
(Geol.  Surv.  Ohio,  11,  pt.  11,  pi.  39,  fig.  3.) 

"A  third  and  larger  specimen  was  found  by  Professor  Newberry  during  the  field  season 
of  1874.  It  includes  an  oblique  view  of  one  side,  and  the  top  of  the  cranium  from  the 
posterior  part  of  the  orbits  to  the  end  of  the  muzzle,  with  the  corresponding  part  of  the 
alveolar  region  of  the  dentary  bone,  with  teeth.  The  bones  of  the  skull  appear  to  have 
been  rather  light,  and  though  the  surface  is  irregular,  the  sculpture  consists  only  of  shallow 
impressions  of  varying  size  and  intervals.  The  orbits  are  also  badly  defined,  but  appear 
to  have  been  large,  and  separated  by  a  narrow  frontal  bone.  The  premaxillary  bone  is 
preserved,  and  shows  clearly  the  sutures  that  separate  it  from  its  fellow  and  from  the 
maxillary.  A  large  foramen — perhaps  the  nostril — separates  it  from  the  maxillary,  so  that 
it  forms  an  irregular  crescent.  It  supports  two  teeth,  of  which  the  anterior  is  the  larger, 
but  there  were  perhaps  others  in  advance,  as  the  alveolar  border  is  imperfect  towards  the 
end  of  the  muzzle.  The  anterior  two  teeth  of  the  maxillary  bone  are  followed  by  a  strong 
groove  which  rises  towards  the  sides  of  the  muzzle.  At  first  sight  this  gives  the  impression 
of  the  maxillo-premaxillary  suture,  and  makes  it  appear  that  both  the  premaxillary  bones 
are  preserved,  and  that  the  foramen  above  described  separates  the  premaxillary  spines, 
instead  of  representing  the  nostril.  The  cutting  edges  of  the  teeth  of  these  bones  have, 
however,  one  direction,  whence  they  represent  one  side  of  the  cranium  only;  were  both 
sides  represented,  the  directions  of  the  tooth  axes  would  be  reversed. 

' '  The  premaxillary  and  maxillary  teeth  exhibit  a  cutting  edge  on  the  outer  posterior 
margin  of  the  distal  half;  the  base  of  the  crown  is  subround  in  section.  The  line-like 
grooves  are  distinct  but  not  numerous,  their  intervals  measuring  75  mm.  Beyond  them 
the  enamel  is  smooth.  The  second  maxillary  tooth  is  larger  than  the  first,  which  is  equal 
to  the  last  premaxillary.  The  third  and  fourth  maxillaries  are  equal  to  the  second,  but 
the  fifth  is  larger  and  longer,  exceeding  all  the  others.  The  teeth  of  the  dentary  bone 
differ  from  those  of  the  upper  jaw  in  having  the  cutting  edge  of  the  crown  on  the  anterior 
aspect,  while  the  posterior  border  is  obtuse.  There  is  an  obtuse  cutting  edge  on  the  pos- 
terior margin  of  the  anterior  mandibular  teeth. 

"This  description  is  derived  from  an  adult  animal,  as  the  maxillary  teeth  in  some 
instances  are  partially  worn  away  by  friction  on  their  anterior  and  outer  faces." 

Measurements. 

mm.  mm. 

Length  of  maxillary  bone  preserved 146       Diameter  of  same  at  base 6 

Length  of  same  supporting  live  teeth 73       Diameter  of  sccondat  base 8 

Length  of  first  maxillary  tooth 15       Length  of  basis  of  fifteen  teeth  of  the  dentary 145 


THE   MICROSAURIAN   FAMILY   SAUROPLEURID.4i. 


169 


i  mjt<  j  a  A   ■  *,•**._ 


Fig.  36. — Mandible  of  Leptophraclus  denlatus  new  species,  from  the 
Linton,  Ohio,  Coal  Measures.  X  I.  Original  in  American  Museum 
of  Natural  History. 


,  Leptophractus  dentatus  new  species. 

Type:  Specimen  No.  1085  G,  American  Museum  of  Natural  History.  Col- 
lected by  Dr.  J.  S.  Newberry. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  type  is  a  single  right  mandible,  nearly  entire,  of  a  rather  large  animal.  The 
specific  characters  for  the  separation  of  the  new  form  from  the  previously  described 
L.  obsoletus  and  L.  lineolatus  are 
the  smaller  size  and  slenderness 
of  the  mandible,  associated  with 
uniform  teeth,  which  are  slender 
and  delicately  fluted. 

There  are  1 7  teeth  preserved, 
the  largest  of  which  is  8  mm. 
in  length.    From  the  posterior 
tooth  the  series  gradually  descends  to  half  this  length  0.5  inch  from  the  anterior 
end  of  the  mandible. 

The  exact  form  of  the  mandible  can  not  be  determined,  but  so  far  as  can  be 
seen  it  is  very  slender,  coming  almost  to  a  point  at  the  anterior  end.  The  poste- 
rior portion  is  wide,  but  apparently  not  very  heavy.  There  is  a  fragment  associated 
with  the  specimen  which  discloses  a  few  teeth,  but  its  position  in  the  cranium  can 
not  be  determined. 

Measurements  of  the  Type  of  Leptophractus  dentatus  Moodie. 

mm.  mm. 

Length  of  mandible,  as  preserved 80  Length  of  most  anterior  tooth 4 . 

Anterior  width  of  mandible 3  Length  of  tenth  posterior  tooth 7 

Posterior  width 28  Width  of  this  tooth  at  base 3 

Leptophractus  lineolatus  Cope. 
Cope,  Proc.  Am.  Phil.  Soc,  xvi,  p.  576,  1877. 

Type:  Specimen  No.  1088  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.    (Plate  22,  fig.  I.) 

Cope's  description  of  this  species,  to  which  I  have  nothing  to  add,  is  as  follows : 

"  This  species  is  based  on  portions  of  the  skull  of  two  individuals  of  large  size.  Both 
upper  and  lower  jaw  with  teeth  are  represented  and  the  teeth  are  of  very  large  size.  The 
deepest  element  preserved  has  been  provisionally  referred  to  the  mandible  and  will  be  so 
described.  This  element  bears  two  types  of  teeth  in  very  heterodont  fashion.  The  teeth  in 
the  back  portion  of  the  jaw  are  rather  short  and  slender.  The  teeth  more  anteriorly  are, 
some  of  them,  very  long,  rather  stout,  with  their  bases  longitudinally  fluted,  as  are  they  all. 
The  longest  tooth  in  the  jaw  measures  slightly  less  than  an  inch.  The  bone  is  not  well 
preserved,  but  seems  to  have  been  ornamented  with  grooves  of  no  great  depth.  The  most 
anterior  teeth  of  the  jaw  are  smaller  than  the  posterior  ones. 

'  The  upper  jaw  is  set  with  teeth  which  are  more  uniform  in  size  and  there  is  but  little 
tendency  to  heterodonty.  These  teeth  are  also  striated  at  their  base  and  all  end  in  a  sharp 
point.  They  are  all,  apparently,  straight.  There  is  no  tendency  to  curve  as  there  is  in  the 
genus  Macrerpeton.  The  upper  teeth  are  more  closely  set  than  are  those  of  the  lower  jaw, 
which  are  rather  distantly  placed. 

"Another  specimen  of  a  smaller  individual  presents  the  same  portions  of  the  skeleton 
and  the  same  characters.  It  is  possible  that  this  skull  will  be  found  to  belong  to  Ichthycan- 
llius  ohiensis  Cope,  which  is  based  on  very  large  vertebras  and  limb  bones.     The  remains 


170 


THE    COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


described  as  Leptophractus  lineolatus  are,  however,  unlike  any  other  skull  remains  which  are 
thus  far  known. 

' '  This  species  represents  one  of  the  largest  types  of  the  Carboniferous  Amphibia  of 
Ohio.  It  probably  attained  a  length  of  several  feet.  It  was  also  the  most  carnivorous  of 
any  of  the  forms." 

Measurements  of  the  Type  Specimen  of  Leptophractus  lineolatus  Cope. 

mm.  mm. 


Length  of  longest  tooth  in  jaw 22 

Anteroposterior  diameter  of  the  same  at  base 6 

Length  of  small  maxillary  tooth 7 

Anteroposterior  diameter  of  same  at  base 2 


Length  of  specimen,  as  preserved 98 

Depth  of  dentary  bone  at  the  middle 30 

Anteroposterior  diameter  of  mandibular  tooth  at 

base 3-5 

Length  of  a  posterior  mandibular  tooth 9 

Collected  by  Dr.  J.  S.  Newberry.     Other  specimens  of  the  species  are  Nos. 
1086  G  and  1087  G,  American  Museum. 

Eurythorax  sublsevis  Cope.* 

Cope,  Proc.  Am.  Phil.  Soc,  p.  177,  1871. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  401,  pi.  xl,  fig.  4,  1875. 

Type:  Specimen  No.  8605  G,  American  Museum  of  Natural  History.     Collec- 
tion of  Dr.  J.  S.  Newberry. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  genus  and  species  were  established  on  a  single,  large,  almost  perfect  inter- 
clavicle  of  peculiar  form.  It  may  or  may  not  be  distinct,  but  it  can  not  be  asso- 
ciated definitely  with  any  known  genus,  so  that 
it  will  be  designated  as  Cope  described  it.  It 
exhibits  on  its  outer  or  lateral  borders  broad, 
smooth  surfaces  for  the  contact  of  the  overlapping 
margins  of  the  lateral  plates.  The  form  is  sub- 
round,  with  a  large  excavation  from  the  posterior 
margin  on  each  side.  The  narrowed  portion  left 
in  the  middle  behind  has  a  convex  outline.  Some 
delicate  radiating  grooves  occur  on  the  exposed 
surface,  but  they  are  very  shallow. 

Measurements  of  Type. 

mm. 

Length  of  interclavicle 71.5 

Greatest  width 78      Fig.  37. — So-called  interclavicle  of  Eurythorax 

Width  of  lateral  concavity 39  sublcevis  Cope.  (Sagenodus.)    X  0.75.    (After 

Cope.) 


*  Dr.  Hussakof  now  regards  (Bull.  Amar.  Mus.  Nat.  Hist.,  vol.  35,  p.  128,  1916)  the  type  specimen  of  this  species 
as  an  opercular  element  of  one  of  the  dipnoan  fishes  [Sagenodus  sublavis  (Cope)].  The  discussion  is  left  here,  however, 
as  a  matter  of  historical  interest. 


CHAPTER  XXII. 

THE   M1CROSAURIAN    FAMILY    ICHTHYCANTHIDyE,    FROM    THE   COAL   MEASURES 

OF  OHIO. 

Family  ICHTHYCANTHIDjE  new  family. 

This  family  is  the  most  recently  recognized  group  of  the  Linton  fauna.  Its 
members,  of  which  there  are  two  known  species,  are  distinct  from  all  other  Coal 
Measures  Amphibia  in  the  possession  of  an  osseous  tarsus  (483,  484),  with  its  asso- 
ciated reptile-like  limb  bones.  There  are  preserved  fine  scutellae  in  a  large  patch 
near  the  vertebral  column.  The  vertebral  spines  are  broad  and  heavy,  with  the 
vertebral  centra  amphiccelous. 

Genus  ICHTHYCANTHUS  Cope,   1877. 

Cope,  Proc.  Am.  Phil.  Soc,  p.  573,  Feb.  3,  1877  (Pal.  Bull.  24). 

Baur,  Beitrage  zur  Morphogenie  des  Carpus  und  Tarsus  der  Vertebraten,  I  Theil,  p.  16,  1888. 

Cope,  Trans.  Am.  Phil.  Soc,  xvi,  p.  289,  fig.  I,  1888. 

Moodie,  Science,  n.s.,  xli,  No.  1044,  p.  34,  1915. 

Moodie,  Am.  Jour.  Sci.,  xxxix,  pp.  509-512,  fig.  2,  May,  1915. 

Type:  Ichthycanthus  ohiensis  Cope. 

The  generic  characters  are  derived  from  the  characters  presented  by  the  pos- 
terior dorsal  and  caudal  vertebrae,  with  adjacent  parts.  The  posterior  limbs  are 
well  developed,  with  distinct  tibia  and  fibula,  osseous  tarsus,  and  5  digits.  Ribs 
elongate,  simple,  curved.  Abdominal  armature  consisting  of  bristle-like  rods  in 
anteriorly  directed  chevrons.  Dorsal  vertebrae  not  elongate,  with  simple  neural 
spines.  Tail  large,  its  vertebrae  ossified,  and  furnished  with  slender  chevron  bones 
which  terminate  in  a  haemal  spine.  Neural  spines  broad  and  directed  backwards ;  the 
caudal  series  somewhat  resembling  that  of  a  fish.     All  the  centra  are  amphiccelous. 

This  genus  differs  from  all  those  with  enlarged  and  sculptured  neural  spines, 
and  from  those  with  abdominal  scutes.  It  is  equally  distinct  from  those  without 
ribs,  abdominal  rods,  or  limbs.  It  is  possible  that  some  of  the  species  referred  to 
Tuditanus,  in  which  these  parts  are  unknown,  may  belong  to  it,  or  that  it  may  be 
established  on  a  small  species  of  Leptophr actus,  a  genus  known  only  as  yet  from  the 
skull.  With  our  present  imperfect  knowledge  of  the  Linton  forms  it  seems  best  to 
refer  /.  ohiensis  and  I.  platypus  to  this  distinct  genus,  Ichthycanthus. 

Ichthycanthus  ohiensis  Cope. 

Cope,  Proc.  Am.  Phil.  Soc,  1877,  p.  573  (Pal.  Bull.  24). 

Type :  Specimen  in  the  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.  Collected  by  Dr.  J.  S. 
Newberry,  in  the  summer  of  1876. 

The  centra  of  the  dorsal  vertebrae  are  about  as  long  as  deep,  and  their  sides  are 
deeply  concave;  there  are  4  anterior  to  the  pelvis  which  are  without  ribs.  The 
caudal  vertebrae  are  robust,  and  7,  from  the  first,  support  a  small  tubercle-like 
diapophysis.  The  chevron  bones  are  short  and  acuminate;  the  neural  spines  are  a 
little  shorter,  narrow,  and  truncate,  and  directed  backwards  at  the  same  angle  as 

171 


172  THE   COAL    MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

the  chevron  bones.     They  are  much  reduced  on  the  eighteenth  caudal  vertebra, 
where  the  chevron  bones  are  considerably  longer. 

The  abdominal  rods  are  quite  slender.  The  hind  limb  is  quite  stout  for  this 
order.  The  femur  is  regularly  expanded  at  both  extremities,  but  the  distal  is  deeply 
and  openly  grooved,  distinguishing  the  condyles,  while  the  proximal  end  is  plane. 
There  is  no  trochanter  visible.  The  ulna  and  radius  are  well  separated,  and  are 
three-fifths  the  length  of  the  femur.  There  is  a  large  fibular  tarsal  bone  of  a  sub- 
quadrate  outline.  In  immediate  contact  with  it  is  probably  the  external  digit  with 
5  phalanges  or  segments ;  the  ungual  is  simply  conic.  The  femur  is  as  long  as  5  dorsal 
vertebrae.  The  ribs  have  expanded,  undivided  heads,  and  extend  to  the  abdominal 
armature. 

Measurements  of  the  Type  of  Ichthycanthus  ohiensis  Cope. 


mm.  mm. 


Length  of  last  10  dorsal  vertebrae 47  Proximal  diameter  of  femur 8 

Length  of  first  23  caudal  vertebrae 117  Width  of  lower  leg 9 

Length  of  a  posterior  rib 29  Length  of  fibula 15 

Length  of  a  posterior  dorsal  vertebra 5  Length  of  tarsal  bone 6 

Length  of  twenty-second  caudal  vertebra 5  Length  of  digit 27 

Length  of  femur 25 

Ichthycanthus  platypus  Cope. 

Cope,  Proc.  Am.  Phil.  Soc.,  pp.  574,  575,  1877. 

Cope,  Trans.  Am.  Phil.  Soc,  xvi,  p.  289,  fig.  1,  1888. 

Baur,  Beitrage  zur  Morphogenie  des  Carpus  und  Tarsus  der  Vertebratcn,  1  Theil,  p.  16,  1888. 

Moodie,  Science,  n.s.,  xli,  No.  1044,  p.  34,  1915. 

Moodie,  Am.  Jour.  Sci.,  xxxix,  pp.  509-512,  fig.  2,  1915. 

Type:  Specimen  No.  7954  G,  and  obverse,  Department  of  Geology,  Columbia 
University.     (Plate  23,  fig.  1.) 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

This  amphibian  is  represented  by  the  same  portions  of  the  skeleton  as  the  preced- 
ing species,  furnishing  a  good  basis  for  comparison.  It  is  very  well  preserved,  display- 
ing the  characters,  especially  of  the  hind  foot,  which  is  almost  entirely  represented. 

Several  features  distinguish  it  from  the  /.  ohiensis,  one  of  which  is  of  more  than 
usual  value  if  correctly  indicated  by  the  fossil.  There  are  10  vertebras  from  the 
anterior  end  to  the  sacrum  preserved  in  place,  and  none  of  them  supports  a  rib, 
nor  are  there  any  ribs  visible  anywhere  on  the  block  of  shale.  I  suspect  that  they 
exist  on  more  anterior  vertebrae,  or  may  have  been  displaced  to  a  more  anterior 
position  than  they  normally  occupy.  The  abdominal  chevrons  are  more  anterior  in 
position  than  are  those  of  the  /.  ohiensis.  The  hind  legs  are  longer  than  in  that 
species;  in  this  one  the  femur  equals  7.5  vertebral  centra  in  length.  The  external 
digit,  on  the  other  hand,  while  bearing  5  phalanges,  is  distinctly  shorter.  The 
fibular  tarsal  is  of  a  transverse  oval,  not  quadrate,  form. 

The  dorsal  centra  are  shorter  and  deeper  than  long;  the  neural  arches  are  ele- 
vated, with  short  but  distinct  zygapophyses,  and  a  flat,  subquadrate,  superiorly 
truncate,  neural  spine.  They  bear  short,  vertically  compressed  diapophyses  near  the 
base  of  the  arches .  The  neural  spines  of  the  caudal  vertebrae  become  rapidly  more  slen- 
der, and  also  diminish  in  length,  while  the  zygapophyses  are  continued  to  the  fifteenth 
vertebra.     The  chevron  bones  are  slender,  and  inclose  a  moderate  haemal  opening. 

The  femur  is  gradually  expanded  to  the  extremities.  Proximally  there  is  a  tro- 
chanteric ala,  besides  the  obtuse  head.    Distally  the  condyles  are  well  distinguished, 


PIATE  2« 


Diplodua 
tooth 


Photograph  of  the  type  specimen  of  I'elion  lyelli  Wyman,  from  the  Ohio  Coal  Measures.  Supposed  to 
represent  the  ancestral  form  of  the  Salientia.     X  2.    Original  in  American  Museum  of  Natural  History. 

Scales  of  Cercariomorphus  parvisquamis  Cope,  a  microsaur  from  the  Ohio  Coal  Measures.  X  10.  Original 
in  American  Museum  of  Natural  History. 

Type  specimen  of  Cercariomorphus  pannsquamis  Cope,  from  the  Ohio  Coal  Measures.  Original  in 
American  Museum  of  Natural  History.     X  0.75. 


THE    MKROSAURIAN    FAMILY    ICHTHYCANTHIDjE.  173 

the  external  or  fibular  being  truncate.  The  fibula  is  less  than  three-fifths  the  length 
of  the  femur,  and  is  expanded  at  both  extremities.  Two  proximal  tarsals  are  dis- 
tinct; the  one  next  the  fibula  is  larger  than  the  other  and  transverse  suboval  in 
form.  It  has  a  median  dividing  ridge  as  though  composed  of  fibulare  and  inter- 
medium coossified.  The  tibiale  is  subtriangular.  There  are  five  distinct  phalan- 
geal tarsals.  The  toes  are,  in  the  order  of  their  lengths,  beginning  with  the  shortest, 
1-2-5-3-4.  Their  phalanges  (including  metatarsals)  are,  in  the  proper  order, 
commencing  with  the  hallux,  3-3-4-5 ?~5,  the  distal  end  of  the  fourth  finger  being 
lost .  These  bones  are  rather  stout,  and  the  unguals  are  simply  conic.  The  form  of 
the  foot  is  short  and  wide.  The  number  of  the  phalanges  is  nearly  similar  to  that 
found  in  Amphibamus  grandiceps,  excepting  that  in  that  species  the  fifth  digit  has 
but  4.  They  are  more  numerous  on  most  of  the  digits  in  Sauropleura  digitata.  Cope 
(Trans.  Am.  Phil.  Soc,  xvi,  289,  fig.  1,  1888)  contributed  the  following  note  on 
Ichthycanthus  platypus: 

"A  reexamination  of  the  type  specimen  of  this  species  from  the  Coal  Measures  of  Ohio, 

preserved  in  the  Museum  of  Columbia  College,  New  York,  enables  me  to  refer  this  species 

to  the  Rachitomi.    The  neural  spines  are  distinct,  showing  that  it  belongs,  probably,  to  the 

Eryopidae.    As  the  skull  is  not  preserved,  I  can  not  determine  the  genus  positively,  but 

refer  it  for  the  present  to  Eryops.    I  append  a  figure  of  the  posterior  foot,  which  displays 

the  characters  of  the  tarsus  of  this  group  for  the  first  time.    The  number  of  tarsals  is  as 

in  a  Theromorph  reptile,  except  that  two  elements  represent  the  cuboid  bone  as  in  the 

reptile,  Stercosternum  tumidum  Cope;  giving  five  elements  in  the  distal  tarsal  row.    There 

is  but  one  centrale  and  no  intermedium.    Two  fragments  of  caudal  vertebra;  adhere  to 

the  specimen." 

Measurements  of  Ichthycanthus  platypus  Cope. 

mm.  mm. 

Length  of  10  dorsal  vertebrae 45  Length  of  fibula 18 

Length  of  15  caudal  vertebrae 55  Diameter  of  fibula  proximally 7 

Length  of  centrum  of  a  dorsal 3.8  Width  of  sole  at  second  row  of  tarsal  bones 17 

Total  elevation  of  a  posterior  dorsal 14  Length  of  foot  to  end  of  third  digit 31 

Length  of  femur 32  Length  of  first  digit 10 

Diameter  of  femur  medially 4.5  Length  of  third  digit 22 

Diameter  of  femur  distally 8.3  Length  of  fifth  digit 20 

The  writer  has  had  the  privilege  of  restudying  this  interesting  specimen  and  has 
already  (484)  described  the  foot  and  tarsus,  as  follows: 

The  only  known  specimen  of  this  anomalous  amphibian  is  incomplete,  repre- 
senting the  posterior  half  of  the  skeleton,  and  an  abundance  of  ventral  scutellas  or  cal- 
cified myocommata.  The  block  of  coal  containing  these  interesting  remains  is  from 
Linton,  Ohio,  and  is  preserved  in  the  geological  collections  of  Columbia  University, 
from  which  institution  Professor  Grabau  very  courteously  forwarded  it  for  study. 

Ichthycanthus  platypus  was  described  by  Cope  from  the  Linton,  Ohio,  Coal 
Measures,  locating  it  doubtfully  in  the  Permian  genus  Eryops  on  account  of  the 
unusual  condition  of  the  tarsus  and  reconsidering  a  former  decision  in  favor  of  a 
Coal  Measures  genus  Ichthycanthus.  In  this  disposition  of  the  species  into  the  Per- 
mian genus  he  is  followed  by  Hay  (317) ;  but  Baur  (28)  regarded  the  form  as  a  mem- 
ber of  the  Coal  Measures  genus  Ichthycanthus,  after  commenting  on  the  later  defi- 
nition by  Cope.  The  type  of  the  genus,  Ichthycanthus,  to  which  Cope  first  allied  the 
species  under  consideration,  is  /.  ohiensis,  a  supposed  amphibian  from  the  Coal 
Measures  of  Linton,  Ohio,  founded  on  incomplete  material. 


174  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

The  form  combines  in  an  unusual  and  remarkable  degree  reptilian  and  amphibian 
characteristics.  The  leg  bones,  pelvis,  and  tarsus  are  all  strikingly  reptilian,  but  the 
phalanges  in  the  arrangement  of  elements  are  so  typically  amphibian  that  if  we  had  no 
other  means  of  diagnosis  we  would  incline  to  locate  this  Coal  Measures  species  among 
the  Amphibia.  The  leg  (plate  23,  fig.  1)  recalls  in  its  structure  that  of  another  Coal 
Measures  species,  Eosauravus  copei  Williston,  which  is,  however,  clearly  a  reptile. 
While  there  is  a  general  degree  of  similarity  between  the  foot  structure  of  Eosauravus 
copei  and  Ichthycanthus  platypus,  yet  there  are  very  great  differences  in  the  phalan- 
geal formula  and  the  arrangement  of  the  tarsal  elements.  These  differences  are  clear 
and  indicate  a  separation  of  the  two  species  into  distinct  classes.  The  phalangeal 
formula  in  the  Eosauravus,  2-3-4-5-4,  is  typically  reptilian;  while  in  the  Ichthy- 
canthus, 2-2-3-3-3,  ^  is  amphibian.  The  tarsus  of  the  Ichthycanthus  is  amphibian 
in  the  presence  of  an  intermedium,  but  this  is  very  small  and  the  remaining  tarsal 
structures  have- nothing  which  might  not  be  found  in  an  early  reptile.  There  may 
be  a  single  or  even  two  centralia  in  the  reptilian  tarsus  among  the  early  forms. 

The  amphibian  nature  of  the  species  having  thus  been  established,  it  remains  to 
give  a  detailed  account  of  its  skeletal  anatomy,  with  comparative  references  to  such 
other  ancient  forms  as  are  available.  Little  can  be  said  of  the  vertebral  column, 
since  only  the  molds  of  a  few  vertebrae  remain,  and  these  are  so  obscured  by  a 
closely  adherent  pellicle  of  carbonaceous  material  that  their  form  can  not  be  dis- 
tinctly discerned.  They  are  high,  with  relatively  broad  neural  spines.  There  are  no 
ribs  preserved.  The  pelvis  is  obscured,  but  it  is  possible  to  determine  the  presence 
of  an  elongate  ilium  and  an  ischium.  The  leg  of  the  left  side  is  the  best  preserved 
of  all  the  elements,  and  it  is  to  this  that  our  attention  will  be  confined.  The  oppo- 
site leg  is  not  so  complete,  yet  all  the  long  bones  and  a  part  of  the  tarsus  are  pre- 
served with  sufficient  clearness  to  corroborate  the  findings  of  the  left  side. 

The  femur,  as  has  been  stated,  is  reptilian  in  appearance.  This  is  due  to  the 
well-rounded  articular  surfaces,  as  though  the  endochondrium  were  well  developed, 
and  to  the  large  development  of  the  greater  and  lesser  trochanters,  which  are  quite 
prominent,  though  these  are  distorted  and  depressed  in  fossilization.  The  bone  is 
stout  and  well  built  and  its  form  suggests  an  active  habit  of  life.  The  tibia  and 
fibula  are  separate,  and  do  not  otherwise  have  sufficiently  noteworthy  characters 
to  call  for  a  special  description  in  this  place,  except  to  note  an  unusual  anterior 
crest  on  the  tibia.  To  the  lower  ends  of  these  bones  articulate  the  first  row  of  tarsal 
elements,  the  tibiale,  intermedium,  and  fibulare.  The  tarsus  is  composed  of  9 
elements  arranged  in  3  rows.  The  proximal  row  is  composed  of  the  tibiale,  the 
intermedium,  and  the  fibulare.  On  the  edge  of  the  tibiale  there  lies  a  portion  of 
one  of  the  caudal  vertebrae,  so  that  the  form  of  this  tarsal  element  is  slightly  ob- 
scured. The  intermedium  is  a  small,  rounded  element  lying  between  the  larger 
elements.  The  fibulare  is  rectangular  and  projects  a  considerable  distance  out  from 
the  tibia,  but  articulates  directly  with  the  large  lateral  distal  tarsal.  The  centrale 
is  triangular  in  form  and  is  opposed  directly  by  the  tibiale  and  tarsalia  1  to  3. 
The  phalanges  are  robust  in  appearance.  The  entire  foot  gives  one  the  impression 
of  a  very  broad  structure.     The  ungual  phalanges  were  apparently  bluntly  clawed. 


CHAPTER  XXIII. 

SUPPOSED  MICROSAURIAN  SPECIES  OF  UNCERTAIN  RELATIONSHIP. 

The  following  three  species  are  so  unusual  and  so  incompletely  known  that  they 
can  not  be  considered  with  any  of  the  above  families:  Brachydectes  newberryi  Cope, 
Linton,  Ohio;  Amblyodon  problematicum  Dawson,  Nova  Scotia;  Proterpeton  gurleyi 
Moodie,  Danville,  Illinois. 

Genus  BRACHYDECTES  Cope,   1868. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  1868,  p.  214. 

Cope,  Trans.  Am.  Phil.  Soc,  1868,  p.  14. 

Cope,  Geol.  Surv.  Ohio,  n,  pt.  11,  p.  388,  1875,  pi.  xxvii,  fig.  2. 

Type:  Brachydectes  newberryi  Cope. 

Cope  (Geol.  Surv.  Ohio,  vol.  n,  pt.  n,  p.  388,  1875),  says: 

"  This  genus  is  indicated  by  two  rami  of  a  mandible  and  a  portion  of  a  premaxillary  only. 
These,  when  compared  with  those  of  (Estocephalus  and  Tuditanus,  from  the  same  locality, 
and  with  others  described  by  authors,  are  so  much  stouter,  i.e.,  shorter  and  more  elevated, 
that  they  evidently  belong  to  a  genus  unlike  either.  The  genus  further  differs  from  (Esto- 
cephalus in  having  the  teeth  of  equal  size  to  the  posterior  part  of  the  series ;  that  is,  to  the 
base  of  the  elevated  coronoid  process.  The  teeth  are  elongate  cylindrical  cones,  with  their 
acute  tips  turned  a  little  posteriorly.  The  fractured  ones  display  a  large  pulp  cavity.  The 
three  premaxillaries  preserved  are  similar,  but  without  curvature  at  the  tips.  They  do  not 
exhibit  striae  or  any  other  sculpture. 

"So  far  as  the  remains  known  go,  the  genus  is  nearer  Hyierpeton  than  any  other. 
According  to  Dawson,  that  genus  is  provided  with  a  large  canine-like  tooth,  at  the  anterior 
extremity  of  the  maxillary,  on  the  inner  row,  which  is  inserted  into  a  distinct  socket.  No 
such  tooth  appears  among  those  of  this  genus.  The  latter  does  not  give  any  indication  of 
the  very  elevated  coronoid  process  of  Brachydectes,  though  the  external  portion  of  the 
dentary  bone  in  that  region  being  lost,  little  can  be  said  about  it." 

Brachydectes  newberryi  Cope. 

Cope,  Proc.  Phila.  Acad.  Nat.  Sci.,  p.  214,  1868. 

Cope,  Trans.  Am.  Phil.  Soc,  1868,  p.  14. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  388,  pi.  xxvii,  fig.  2,  1875. 

Type :  Specimen  No.  8604  G,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton   Ohio,  Coal  Measures. 

Cope  (Geol.  Surv.  Ohio,  vol.  n,  pt  II,  p.  388,  1875)  says  of  this  form: 

"  The  species  is  represented  by  one  nearly  perfect  ramus  mandibuli,  one  dentary  bone 
and  one  premaxillary,  probably  not  complete. 

' '  The  dentary  bone  appears  to  have  been  attached  by  suture  to  the  articular  and  angular, 
as  its  free  margin  has  very  much  of  the  outline  of  that  suture  in  Amphiuma  and  lizards.  The 
coronoid  process  would  also  seem  to  be  a  part  of  the  same  bone  as  in  Amphiuma  and  Meno- 
pcma,  and  not  composed  of  the  coronoid  bone  as  in  lizards.  It  rises  immediately  behind  the 
last  tooth,  and  displays  no  suture. 

175 


I76  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

"The  lower  portion  of  the  dentary  is  prolonged  into  an  acute  angle.  This  is  separated 
by  a  deep  and  wide  concavity  from  the  superior  posterior  prolongation,  which  is  obtuse, 
and  rises  at  once  into  the  coronoid  process.  Teeth  on  this  dentary,  seven ;  the  same  number 
is  on  the  preserved  ramus ;  this  number  is  suspected  to  be  complete,  or  nearly  so.  The  teeth 
terminate  at  the  obvious  termination  of  each  ramus,  which  is,  it  is  true,  slightly  obscured. 
These  teeth  are  the  longest  in  the  Microsauria  in  relation  to  the  depth  of  the  ramus,  equal- 
ing the  largest  in  (Estocephalus.  They  are  doubtless  exposed,  as  are  some  of  those  of  the 
last-named  genus,  by  the  splitting  away  of  the  outer  parapet  of  the  dentary  bone.  As  no 
traces  of  alveoli  have  been  thus  rendered  visible  I  suspect  the  dentition  to  have  been 
acrodont,  as  in  some  existing  Batrachia. 

' '  No  external  surface  of  the  mandible  remains,  but  there  are  no  impressions  of  sculpture 
on  the  matrix.    A  little  external  face  of  the  premaxillary  displays  none. 

"  The  species  is  dedicated  to  Professor  John  S.  Newberry,  the  able  director  of  the  Geo- 
logical Survey  of  Ohio,  and  discoverer  of  most  of  the  Batrachia  herein  described." 

Measurements  of  the  Type. 

mm.  mm. 

Length  of  ramus  of  mandible  (imperfect) 22  Length  of  dentary 16 

Depth  at  last  tooth 5  Depth  at  coronoid  process 7.5 

Length  of  exposed  tooth 3.5      Depth  at  first  tooth 3 

Genus  PROTERPETON  new  genus. 

Type:  Proterpeton  gurleyi  Moodie. 

Known  from  a  single  vertebra.  Spine  very  high  and  heavy,  the  neural  canal 
large. 

Proterpeton  gurleyi  new  species. 

Type:  Specimen  No.  13,296,  Walker  Museum,  University  of  Chicago. 

Horizon  and  locality:  Coal  Measures  near  Danville,  Illinois. 

The  vertebra,  as  preserved,  is  well  characterized  by  the  figure  (plate  22,  fig.  2). 
The  spine  is  high  and  heavy,  the  neural  canal  is  large,  and  the  centrum  reduced. 
The  form  is  very  unusual.  It  is  apparently  from  the  cervical  region,  as  there  are  no 
indications  of  zygapophyses,  transverse  processes,  or  haemal  arches,  although  they 
may  have  been  abraded;  apparently  not,  however.  The  type  specimen  was  dis- 
covered near  Danville,  Illinois,  about  the  horizon  of  the  Danville  coal,  so  that  it  is 
quite  high  in  the  Allegheny  series  of  the  Pennsylvanian  and  of  about  the  same 
horizon  as  the  phalangeal  bone  from  Breeze,  Illinois,  which  may  be  provisionally 
associated  with  this  form.  There  is  no  assurance  that  Proterpeton  gurleyi  is  an 
amphibian.     The  vertebra  may  have  belonged  to  a  fish. 

Measurements  of  the  Type  of  Proterpeton  gurleyi  Moodie. 

mm.  mm. 

Entire  height  of  vertebra 24  Height  of  neural  canal  (crushed  ?) 6.5 

Width  at  side  of  neural  canal 21.5  Width  of  vertebral  centrum  anteroposteriorly 5.5 

Width  of  neural  canal 13  Height  of  neural  spine  from  top  of  neural  canal  ...  9 

Genus  AMBLYODON  Dawson,   1882. 
Dawson,  Phil.  Trans.  Roy.  Soc.  London,  pt.  11,  p.  644,  pi.  40,  figs.  57-61,  1882. 

Type:  Amblyodon  problematicum  Dawson. 

This  genus  was  described  by  Dawson  in  1882  from  very  imperfect  remains.  He 
says  that  it  is  "characterized  by  stout  cylindrical  teeth,  blunt  at  the  apices;  but 
otherwise  imperfectly  known." 


SUPPOSED   MICROSAURIAN   SPECIES   OF   UNCERTAIN   RELATIONSHIP. 


177 


Amblyodon  problematicura  Dawson. 
Dawson,  Phil.  Trans.  Roy.  Soc.  London,  pt.  u,  p.  644,  pi.  40,  figs.  57  to  61,  1882. 

TyPe:  Specimen  No.  3061-10,  Peter  Redpath  Museum,  McGill  University- 
Horizon  and  locality:  Coal  Measures  of  Nova 

Scotia. 

A  fragment  of  a  jaw  1  cm.  in  length  has  10 

cylindrical  teeth,  simple  and  smooth,  with  large 

pulp  cavities  and  rounded  regularly  at  the  apices. 

With  these  are  4  vertebrae  of  the  usual  type, 

measuring  together  1  cm.    Fragments  of  cranial 

bones  also  occur  and  are  obscurely  pitted.    There 

is  also  what  seems  to  be  the  shaft  of  a  limb 

bone  and  a  few  oval  scales.      A  flat  and  some- 
what rhombic  bone,  with  a  style  at  one  side, 

may  possibly  be  a  thoracic  plate  or  possibly  a 

parasphenoid. 

The  material  is  too  scanty  for  any  satisfactory  description  of  this  animal,  but  it 

is  provisionally  named  Amblyodon  problematic  urn. 


Fig.  38. — Skeletal  elements  of  Amblyodon  sp. 
from  the  Coal  Measures  of  Nova  Scotia, 
a,  tooth,  X  25;  b,  section  of  tooth,  X  25; 
d,  fragment  of  thoracic  plate;  /,  shaft  of 
limb  bone;  e,  rib. 


CHAPTER  XXIV. 

THE  TEMNOSPONDYLOUS  AMPHIBIA  OF  THE  COAL  MEASURES 
OF  NORTH  AMERICA. 

DEFINITION  OF  THE  ORDER  TEMNOSPONDYLIA,  ZITTEL,  1887. 

Zittel,  Handbuch  der  Paleontologie,  Abth.  I,  Bd.  3,  p.  384,  1887. 

Terrestrial  or  semi-aquatic  vertebrata;  skull  bones  pitted  and  grooved ;  lateral- 
line  canals  present  in  well-developed  form;  pineal  foramen  sometimes  absent; 
sclerotic  plates  present ;  vertebras  rachitomous  or  embolomerous ;  notochord  partly 
persistent;  one  or  two  sacral  vertebrae;  tail  present,  long  or  short;  limbs  and 
girdles  well  developed;  limb  bones  well  ossified  and  bones  of  arm  and  leg  sepa- 
rate; pectoral  and  pelvic  girdles  composed  of  the  usual  stegocephalian  elements; 
an  osseous  pubis  present;  a  cleithrum  present  on  the  scapula;  carpus  and  tarsus 
ossified,  carpals  11  and  tarsals  12  in  one  form;  phalangeal  formula,  2,  3,  3,  4,  2 
for  the  hand  and  2,  3,  3,  3,  2  for  the  foot;  fore  and  hind  limbs  pentadactyl  in  a 
few  forms;  venter  covered  with  an  armature  of  osseous  scutes,  sometimes  over- 
lapping; skin  of  back  bare  or  armored  with  heavy  plates;  ribs  heavy,  double- 
headed,  curved  and  moderately  long,  or  short ;  body  short  and  heavy,  as  compared 
to  skull  about  2  to  1 . 

Range:  Coal  Measures  to  upper  Permian. 

Distribution:  North  America:  Illinois,  Kansas,  Oklahoma,  Texas,  and  Penn- 
sylvania; Europe:  Germany,  Bohemia;  France;  Asia:  India. 

Family  CRICOTID^  Cope,  1884. 

Cope,  Am.  Nat,,  xvm,  p.  38,  1884. 

General  form  of  the  body  elongate,  with  triangular  skull  and  short,  stout 
limbs.  Snout  narrow,  orbits  large,  elongated  oval,  situated  near  the  middle  of 
the  skull.  External  bones  faintly  sculptured,  sensor}-  canals  conspicuous,  parietal 
foramen  large.  Teeth  conical,  of  unequal  size.  Presacral  vertebras  composed  of 
horseshoe-shaped  pleurocentra  and  hypocentra,  the  former  alone  supporting  the 
neural  arch.  In  the  caudals  the  pleurocentra  and  hypocentra  form  complete 
rings,  and  both  elements  take  part  in  the  support  of  the  neural  arch,  but  the 
haemal  arch  is  borne  exclusively  by  the  hypocentra.  A  close  abdominal  armor  of 
imbricate  scales,  arranged  in  a  chevron  pattern.  Caudal  vertebrae  numerous. 
Chevrons  cobssified  with  the  intercentra. 

Genus  SPONDYLERPETON  Moodie. 

Moodie,  Kans.  Univ.  Sci.  Bull.,  vi,  No.  2,  p.  355,  1912. 

Type:  Spondylerpeton  spinatum  Moodie. 

The  genus  is  based  on  a  specimen  consisting  of  9  imperfect  vertebrae,  from  the 
caudal  region  of  a  relatively  large  amphibian.  The  present  genus  exceeds  Diplo- 
spondylus  from  the  Gaskohle  of  Bohemia  (251)  by  twice  its  size  and  is  about  two- 
thirds  the  size  of  Cricotus  heteroclitus  Cope  (98)  from  the  Permian  of  Kansas.  The 
vertebras  are  twice  as  high  as  wide,  differing  thus  from  Cricotus,  in  which  the  ver- 
tebras are  nearly  circular.  A  character  which  is  of  great  importance  is  the  large  size 
of  the  intercentrum,  which  almost  equals  the  pleurocentrum  in  size.  It  is  similar 
to  the  pleurocentrum  in  shape,  except  for  the  attached  neurocentrum  and  chevron  on 
the  latter.    The  present  genus  differs  from  Diplospondylus  in  the  greater  length  of 

178 


THE   TEMNOSPONDYLOUS    AMPHIBIA. 


179 


the  intercentrum  and  pleurocentrum,  in  the  greater  size,  in  the  larger  proportions 
of  the  neurocentrum,  and  the  greater  proportionate  size  of  the  intercentra. 

Spondylerpeton  spinatum  Moodie. 
Moodie,  Kans.  Univ.  Sei.  Bull.,  vi,  No.  2,  pp.  355-357,  pi.  8,  figs.  1  and  2;  pi.  9,  fig.  1,  1912. 

Type :  Specimen  No.  793  (26)  and  obverse,  Yale  University  Museum. 

Horizon  and  locality:  Mazon  Creek,  near  Morris,  Illinois. 

The  species  is  very  imperfectly  known.  Sufficient  is  present,  however,  to  show 
its  wide  generic  differences  from  other  forms  of  the  Cricotidae.  These  characters 
are  of  a  phylogenetic  nature  and  indicate  the  more  primitive  nature  of  the  present 
form,  as  we  would  expect  from  its  geological  position.  The  sutures  separating  the 
four  vertebral  elements  are  clearly  ap- 
parent. The  pleurocentral-neurocentral 
suture  is  apparent  in  4  vertebras. 

There  is  but  a  single  pleurocentrum 
preserved  complete.  This  shows  the 
form  of  the  attached  neurocentrum  and 
chevron,  which  corresponds  to  the  hypo- 
centrum  pleurale  according  to  Fritsch. 
The  pleurocentrum  is  flattened  laterally, 
with  a  rather  large  canal  for  the  noto- 
chord.  Its  sides  are  marked  with  4 
longitudinal  grooves.  Surfaces  for  the 
attachment  of  the  ribs  are  not  present, 
and  for  this  reason,  as  well  as  the  pres- 
ence of  chevrons,  the  vertebras  are  sup- 
posed to  be  caudals.  As  such  they  rep- 
resent an  animal  of  some  3  or  4  feet  in 
length.  It  was  the  giant  of  the  Mazon 
Creek  Amphibia.     (Plate  4,  figs.  1,  2.) 

Attached  to  the  upper  side  of  the 
pleurocentrum  by  a  sutural  union  occurs 
the  neurocentrum.  The  neural  arch  is  quite  large  and  is  oval  in  outline,  although 
somewhat  constricted  at  the  tip.  The  spine  of  the  neurocentrum  is  rather  long  and 
broad  at  the  base,  measuring  12  mm.  across  the  anterior  zygapophysis.  The  neuro- 
centrum is  laterally  flattened  and  ends  in  a  rather  acute  and  somewhat  rugose 
point.  It  was  probably  tipped  with  cartilage.  The  anterior  zygapophysis  occurs 
well  down  on  the  neurocentrum,  its  lower  edge  being  5  mm.  from  the  suture  sepa- 
rating the  pleurocentrum  and  the  neurocentrum.  The  posterior  zygapophysis  occurs 
quite  high  up  on  the  neurocentrum  and  lies  at  a  distance  of  15  mm.  from  the  pleuro- 
neurocentral  suture,  thus  indicating  an  extreme  posterior  inclination  of  the  neural 
spine.  The  posterior  zygapophysis  of  the  best  preserved  vertebra  is  separated 
from  its  mate,  the  anterior  zygapophysis,  in  the  next  succeeding  vertebra  by  a 
space  of  5  mm. 


Fig.  39. — The  vertebrae  of  Spondylerpeton  spinatum 
Moodie,  the  only  known  temnospondyle  from  the 
Mazon  Creek  shales.  X  I.  hy,  hypocentrum;  inc, 
intercentrum;  pc,  pleurocentrum;  nc,  neurocentrum. 


180  THE    COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

The  ventral  surface  of  the  pleurocentrum  bears  a  structure,  which  is,  without 
doubt,  a  chevron,  although  the  character  of  its  opening  can  not  be  determined.  It 
is  elongated  and  is  united  by  a  broad  base  to  the  pleurocentrum.  Its  union  is  by  a 
clearly  defined  suture,  which  is  apparent  in  3  vertebrae.  The  condition  represented 
by  the  specimen  duplicates  almost  exactly  the  condition  figured  by  Cope  for  the 
caudal  region  of  Cricotus  Cope.* 

The  intercentrum  of  the  present  form  is  fully  as  large  as  the  pleurocentrum. 
The  significance  of  this  has  already  been  mentioned.  The  body  of  the  centrum  is 
pierced  by  a  large  notochordal  canal. 

Measurements  of  the  Type  of  Spon'dyi.erpeton  spinatum  Moodie. 

mm.  mm. 

Length  of  specimen 60  Height  of  intercentrum 10.5 

Length  of  pleurocentrum 1 1 .5  Height  of  chevron 3 

Height  of  pleurocentrum  to  base  of  neurocentrum .  20  Length  of  chevron 18 

Length  of  neurocentrum 33  Width  of  notochordal  opening  in  centrum 5 

Width  of  neurocentrum  at  base 9  Height  of  same 4.5 

Width  across  anterior  zygapophysis 12  Height  of  neural  canal 12 

Width  across  posterior  zygapophysis 10  Greatest  width  of  neural  canal 6 

Length  of  intercentrum 10 

Family  ERYOPIDiE  Cope,  1882. 

Cope,  Am.  Nat.,  xvi,  p.  334,  1882. 

Large,  terrestrial  or  amphibious  vertebrata;  skull  bones  deeply  marked  with 
pits  and  grooves  which  take  the  form  of  lateral-line  canals;  infra-  and  supra- 
orbital canals,  antorbital  commissure,  jugal  canal,  and  occipital  cross-commissure 
of  the  lateral-line  system  present  in  Eryops  megacephalus  Cope ;  carpus  and  tarsus 
osseous;  pubis  an  osseous  plate,  surrounded  in  life  by  a  large  amount  of  cartilage; 
fore  and  hind  limbs  pentadactyl ;  orbits,  in  the  typical  genus,  located  far  back  on 
the  skull  and  near  the  median  line ;  cleithrum  present  on  the  scapula ;  vertebrae 
rachitomous,  the  intercentrum  supporting  the  arch  in  the  dorsal  region ;  para- 
sphenoid  well-developed  or  reduced;  teeth  on  pterygoids,  palatines,  prevomers, 
and  parasphenoid. 

Range:  Upper  Pennsylvanian  to  Permian. 
Distribution:  America,  Europe,  Asia. 

Genus  ERYOPS  Cope,  1877. 

Type:  Eryops  megacephalus  Cope. 

Skull  long,  comparatively  narrow;  proportion  of  length  to  breadth  about  9 
to  7.  Roof  bones  coarsely  sculptured  posteriorly,  finely  sculptured  anteriorly. 
Nasals  and  premaxillae  very  large;  frontals  excluded  from  the  orbits  by  junction 
of  pre-  and  post-frontals.  Pterygoids  not  meeting  in  the  median  line ;  parasphe- 
noid dagger-shaped,  tapering  gradually  to  a  point  just  in  front  of  the  palatine 
foramen;  prevomers  large.  Orbits  subcircular,  situated  in  the  posterior  half  of  the 
skull;  nares  subovate,  remote,  at  a  considerable  distance  from  the  tip  of  the  skull. 
Many  minute  denticles,  on  pterygoids,  palatines,  prevomers,  and  parasphenoid. 
Teeth  circular  in  cross-section,  strongly  ribbed  near  the  base,  dentine  strongly 
infolded.  Three  large  teeth  on  each  palatine.  Mandible  without  postcotyloid 
process.  Vertebras  rachitomous.  Ribs  double-headed.  Pubis  osseous.  Three 
species  (Permian),  E.  megacephalus,  E.  lotus,  and  E.  willistoni,  are  assigned  to 
the  genus. 

Eryops  is  represented  in  the  Carboniferous  deposits  of  North  America  by  incom- 
plete vertebral  remains  described  by  Case  (94)  from  near  Pittsburgh,  Pennsylvania. 

*Cope,  E.  D.,  Trans.  Am.  Phil.  Soc,  xvi,  p.  246,  1890. 


MOODIE 


PLATE  25 


■\>, 


i.& 


'/ 


v->: 


&&tfgU^ 


-^ 


1.  Photograph  of  type  specimen  of  Erpetosaurus  (Tiit/itanits)  nu/inlns  Cope,  from  the  Coal  Measures  of 

Linton,  Ohio.     X  1.3.     Original  in  American  Museum  of  Natural  History. 

2.  Photograph  of  type  specimen  of  Erpetosaurus  tabulalus  Cope,  from  the   Coal    Measures   of   Linton, 

Ohio.     X  2.     Original  in  the  Zoological  Collections  of  Columbia  University. 

3.  Photograph  of  the  impression  of  Stegops  divaricate  Cope,  from  the  Coal  Measures  of  Linton,  Ohio. 

X  2.     The  specimen  figured  is  in  the  American  Museum  of  Natural  History.     Its  obverse   is   in 
the  collections  at  Walker  Museum,  I'niversitv  of  Chicago. 

4.  Type  and  only  known  specimen  of  .Vi<  rrrpr/on  caiufatittH    Moodie,    a  branchiosaur   from    the    Coal 

Measures  shales  of  MilOB  Creek,  Illinois.     X  2.     Original  in   collections   at    Walker   Museum, 
University  of  Chicago, 


THE   TEMNOSPONDYLOUS    AMPHIBIA.  l8l 

Eryops  sp.  indet.  Case,  1908. 
Case,  Annals  Carnegie  Mus.,  iv,p.  234,  pi.  59,  1908. 

A  dorsal  vertebra  is  very  probably  from  this  genus.  The  specimen  consists  of  a 
nearly  perfect  vertebra,  lacking  only  the  anterior  zygapophysis  and  the  upper  por- 
tion of  the  neural  spine  (plate  i8,  fig.  2).  It  shows  no  character  that  would  war- 
rant its  separation  from  the  genus,  and  indicates  a  medium-sized  individual.  The 
zygapophyses  have  clean-cut  articular  faces.  The  pleurocentra  are  thickened  above, 
with  just  well-defined  articular  faces,  which  were  applied  to  faces  on  the  neural  arch 
posterior  to  the  origin  of  the  transverse  process.  The  intercentrum  is  of  the  familiar 
half  moon-shape,  thick  and  heavy  below,  and  thinner  toward  the  extremities;  the 
anterior  edge  is  marked  near  the  top  by  the  indentation  found  on  the  intercentra  of 
Eryops. 

Height  of  the  vertebra  from  the  middle  of  the  lower  face  of  the  intercentrum  to 
the  middle  of  the  neural  canal,  0.035  m- ;  width  of  intercentrum  0.026  m. 

The  second  recognizable  specimen  is  a  neural  spine  from  the  caudal  series.  This 
is  without  question  a  portion  of  the  skeleton  of  an  Eryops.  Similar  spines  were 
described  by  Cope  as  Eryops  (Epicordylus)  erythrolithicus,  but  later  discoveries 
seem  to  show  that  similar  characters  occur  in  other  species  of  the  genus  as  well.  The 
apex  of  the  spine  is  bifurcate;  the  space  between  extremities  is  concave  and  per- 
fectly smooth ;  below  the  sides  of  the  spine  are  rather  rugose  and  marked  with  ridges. 
The  lower  portion  of  the  spine  is  elongated  anteroposteriorly  and  the  edges  are 
marked  with  sharp,  double  ridges. 

Three  ribs  also  belong,  in  all  probability,  to  the  genus  Eryops.  The  head  of 
each  rib  is  broad  and  the  articular  edge  is  divided  between  two  faces  which  meet  at 
an  angle  somewhat  greater  than  a  right  angle;  the  two  faces  are  continuous.  The 
shaft  is  somewhat  flattened  and  in  the  undistorted  specimens  is  gently  curved.  The 
length  of  the  largest  rib  is  about  0.07  m. 

Other  than  these  specimens  there  are  several  small  intercentra  (94)  and  the 
neural  spine  of  a  caudal  vertebra  from  some  undetermined  amphibian. 

Family  MACRERPETIDjE  Moodie,  1909. 

Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  art.  xxv,  p.  354,  pi.  lix,  fig.  I,  1909. 

It  has  seemed  necessary  to  propose  a  new  family  for  the  reception  of  the  single 
species  Macrerpeton  (Tuditanus)  huxleyi  Cope.  The  characters  exhibited  by  this 
species  are  so  different  from  those  offered  by  other  members  of  the  Carboniferous 
Microsauria  that  it  is  clearly  distinct.  In  its  cranial  characters  and  the  position  of 
the  orbits  it  approaches  most  nearly  to  Eryops  megacephalus  Cope  from  the  Per- 
mian of  Texas.  In  some  of  its  characters  the  present  form  shows  a  similarity  to 
Dasyceps  bucklandi  Lloyd  (324),  from  the  Permian  of  Kenilworth,  England;  more 
especially  is  this  similarity  found  in  the  form  of  the  skull,  the  size  and  shape  of  the 
teeth,  and  the  posterior  position  of  the  orbits,  and  their  wide  removal  from  the 
border  of  the  skull.  Only  a  fragment  of  the  skull  has  hitherto  been  known,  but  re- 
peated study  of  this  fragment  (123)  has  disclosed  the  wide  diversity  (462)  of  its 
characters.    An  almost  complete  skull,  described  below,  substantiates  the  charac- 


1 82  THE   COAL    MEASURES   AMPHIBIA    OF    NORTH    AMERICA. 

ters  based  on  the  fragment.    Another  species  is  here  added  to  the  genus,  based  on  a 
portion  of  a  mandible  and  a  portion  of  the  skull. 

The  family,  Macrerpetidae,  may  be  defined  (465)  as  follows : 

Skull  larger  than  in  any  other  known  microsaurian,  unless  Baphetes  proves 
to  be  microsaurian;  cranial  elements  sculptured  with  pits  and  coarse  grooves; 
lacrimal  element  present,  teeth  large,  curved  inwards  and  fluted;  mandible 
heavy;  orbits  located  far  back  on  the  skull  and  near  the  median  line  so  that  the 
interorbital  space  is  about  half  the  space  from  the  outer  edge  of  the  orbit  to  the 
border  of  the  skull,  thus  approaching  the  condition  known  in  Eryops;  the  ribs  (?) 
are  strong,  heavy,  and  curved,  with  an  incipient  tubercle. 

Genus  MACRERPETON  Moodie,  1909. 
Moodie,  Jour.  Geol.,  xvn,  No.  I,  pp.  72-74,  fig.  17,  1909. 

Type:  Macrerpeton  huxleyi  Cope. 

The  genus  Macrerpeton  was  proposed  for  the  reception  of  the  amphibian  species 
described  by  Cope  as  Tuditanus  huxleyi  (123).  This  form  he  placed  provisionally  in 
genus  Tuditanus,  since  it  seemed  to  present  the  same  type  of  sculpturing  of  the 
cranial  elements  similar  to  that  found  in  T.  radiatus  Cope.  But  this  species  has 
been  removed  from  Tuditanus  and  placed  in  a  new  genus,  Erpetosaurus  (462).  A 
close  study  of  the  type  specimen  of  Tuditanus  huxleyi  Cope  shows  (465)  great 
variation  and  distinction  from  any  of  the  species  described  from  Linton,  Ohio,  or 
indeed  from  any  Carboniferous  form  thus  far  known. 

The  specimen  represents  the  left  side  of  the  face,  and  the  characters  exhibited  by 
the  fragment  are  supported  by  more  complete  material  (No.  2933,  Am.  Mus.  Nat. 
Hist.).  The  skull  shows  a  close  approach  to  the  higher  labyrinthodonts  in  its  shape. 
The  orbits  are  far  removed  from  the  border  of  the  skull.  The  arrangement  of  the 
bones  of  the  skull  resembles  that  of  Capitosaurus  from  the  Keuper  of  Europe.  The 
jaw  is  for  the  most  part  slender,  with  a  pronounced  downward  inflection  at  the  coro- 
noidal  part.  The  teeth  are  heavy  and  strong  and  are  curved  backwards.  They 
have  the  strong  longitudinal  fluting  which  is  characteristic  of  the  labyrinthodonts. 
Another  character  which  is  distinctive  is  the  pattern  of  cranial  sculpture.  This 
consists  of  inosculating  pits  and  grooves  of  a  coarse  character  and  compares  favor- 
ably with  the  sculpturing  of  Triassic  labyrinthodonts.  If  Macrerpeton  really  rep- 
resents a  labyrinthodont  form  of  Amphibia  it  is  the  oldest  of  the  known  Labyrintho- 
dontidae,  since  it  seems  probable  that  the  Eosaurus  vertebrae  came  from  a  higher 
horizon. 

Macrepeton  huxleyi  Cope,  1874. 

Cope,  Trans.  Am.  Phil.  Soc,  xv,  p.  274,  1874. 

Cope,  Geol.  Surv.  Ohio,  11,  pt.  11,  p.  397,  pi.  xxxiv,  fig.  2,  1875. 

Lesley,  Dictionary  of  Fossils,  p.  1237,  1890. 

Moodie,  Jour.  Geol.,  xvn,  No.  1,  p.  72,  fig.  17,  1909. 

Moodie,  Bull.  Am.  Mus.  Nat.  Hist.,  xxvi,  art.  xxv,  p.  354,  pi.  lix,  fig.  1,  1909. 

Type:  Specimen  No.  119,  American  Museum  of  Natural  History.  Collection  of 
Dr.  J.  S.  Newberry.    (Plate  26,  fig.  2.) 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures. 

The  first  part  of  the  following  account  of  the  species,  Macrerpeton  huxleyi 
Cope,  is  a  quotation  of  Cope's  description  (123)  of  the  type  specimen,  and  the  second 
part  deals  with  the  description  of  the  new  material.    Cope  says  the  species  is — 


MOOOII 


Pl.AT£   26 


2 


1.  Photograph  of  type  specimen  of  Erpetosaurus  tuberculatus  Moodie,  from  the  Ohio  Coal 
Measures.     X  1.     Original  in  American  Museum  of  Natural  History. 

Z.  Photograph  of  type  of  Macrerpeton  huxleyi  Cope,  from  the  Ohio  Coal  Measures.  X  1. 
Original  in  American  Museum  of  Natural   History. 


THE    TEMNOSPONDYLOUS    AMPHIBIA.  1 83 

"  Represented  by  a  considerable  portion  of  the  face  and  muzzle  of  a  single  individual. 
A  portion  of  the  left  mandible,  supporting  three  teeth,  remains  in  place,  and  almost  the 
entire  boundary  of  the  right  orbit  is  preserved. 

'  The  fragment  indicates  a  much  larger  species  than  any  other  referred  to  the  genus, 
and,  next  to  the  Le ptophractus  obsoletus,  the  largest  of  the  Batrachians  of  the  Ohio  Coal 
Measures.  Without  more  complete  remains,  it  is  not  easy  to  determine  its  generic  relations 
finally. 

"  The  form  of  the  head  is  probably  elongate,  and  the  muzzle  neither  very  obtuse  nor 
elongate.  The  orbit  is  rather  small,  and  near  the  middle  of  the  length  of  the  specimen, 
which  is,  however,  incomplete  at  both  ends.  The  sculpture  of  the  surface  of  the  head  pos- 
terior to  the  orbits,  as  well  as  round  their  borders  and  for  some  distance  in  front  of  them, 
consists  of  a  rather  coarse  pitting.  On  the  middle  line,  between  the  orbits  and  on  the  muz- 
zle, the  intervals  become  narrower,  and  are  confluent  into  transverse  ridges  or  a  delicate 
reticulation.    The  surface  of  the  mandible  displays  a  coarse  reticulation. 

"  The  teeth  are  stoutly  conic,  and  with  delicately  striate  grooved  cementum.  They  are 
slightly  recurved. 

"  This  species  differs  from  the  T.  radiatus  and  T.  obtusus  in  the  absence  of  the  area  into 
which  the  sculpture  is  thrown. 

"  Longitudinal  diameter  of  orbit,  19  mm.;  length  of  alveolar  border  supporting  three 
teeth,  13  mm. ;  diameter  of  base  of  tooth,  3  mm. ;  eight  pits  in  10  mm. 

"  Dedicated  to  Professor  T.  H.  Huxley,  facile  princeps  among  English  systematists,  and 
an  important  contributor  to  the  knowledge  of  the  extinct  Batrachia." 

The  following  discussion  of  the  cranial  elements,  based  on  the  writer's  studies 
(462,  465)  of  the  type,  may  be  appended  to  Professor  Cope's  original  description. 
The  sutures  bounding  a  few  of  the  elements  have  been  made  out  in  part.  The 
prefrontal  element  seems  well  assured.  It  lies  well  in  front  of  the  orbit,  much  as  in 
the  skull  of  Capitosaurus  from  the  Keuper  of  Europe.  The  lacrimal  is,  apparently, 
a  very  large  bone,  though  its  entire  extent  is  not  assured. 

The  maxilla  is  a  long,  narrow  element  on  the  border  of  the  skull.  The  suture 
separating  this  from  the  lacrimal  and  jugal  is  quite  clear.  The  teeth  which  the 
maxilla  undoubtedly  bore  are  hidden  by  the  remains  of  the  mandible,  which  lies 
partly  on  the  edge  of  the  skull.  The  jugal  is  a  very  large  element  and  its  boundaries 
seem  well  assured.  Its  size  and  relations  recall  the  condition  in  Capitosaurus.  It 
forms  a  part  of  the  external  boundary  of  the  orbit.  The  lateral  suture  of  the  post- 
orbital  is  evident  and  is,  as  shown  in  the  figure,  somewhat  curved.  The  remaining 
elements  preserved  on  the  fragmentary  skull  can  not  be  accurately  determined, 
though  their  probable  position  is  indicated  in  plate  30,  fig.  2,  the  lettering  being 
based  on  the  arrangement  of  these  elements  in  Capitosaurus. 

The  lower  jaw  is  poorly  preserved,  but  what  remains  shows  evidence  of  being 
sculptured  somewhat  after  the  manner  of  the  cranial  elements.  It  bore  strong 
recurved  teeth  which  are  longitudinally  striate. 

Measurements  of  the  Type  Specimen  of  Macrerpeton  huxleyi  Cope. 

mm.  mm. 

Length  of  portion  preserved 120  Length  of  jaw,  as  preserved 75 

Maximum  width  of  specimen 58  Width  of  jaw  at  widest  part 11 

Length  of  orbit 20  Length  of  longest  tooth 8 

Width  of  orbit 14  Width  of  tooth  at  base 4 


1 84 


THE    COAL    MEASURES   AMPHIBIA    OF    NORTH    AMERICA. 


DESCRIPTION  OF  ADDITIONAL   MATERIAL  OF  MACRERPETON  HUXLEYI. 

The  additional  material  of  this  species  which  has  come  to  hand  consists  of  an 
almost  complete  skull  (American  Museum  No.  2933,  two  portions);  another  frag- 
mentary skull  (American  Museum  No.  8572  G  and  8532  G);  a  portion  of  an 
interclavicle  (American  Museum  No.  8006) ;  two  incomplete  vertebrae  (American 
Museum  No.  8007) ;  and  another  fragmentary  element  possibly  representing  a 
scapula  of  this  species  (American  Museum  No.  8008). 

The  skull  has  essentially  the  shape  outlined  (462)  from  a  study  of  the  fragmen- 
tary type  specimen.  The  muzzle  was  drawn  slightly  too  broad,  but  otherwise  the 
restoration  is  fairly  accurate.  The  specimen  is  distorted  and  imperfect,  but  enough 
is  preserved  to  give  a  good  idea  of  the  shape  and  something  of  the  structure  of  the 
skull.  A  portion  of  the  obverse  is  preserved.  The  back  part  of  the  skull  is  broken, 
so  that  the  occiput  can  not  be  studied. 

The  length  of  the  skull  is  one  and  two-fifths  the  greatest  breadth  (across  the 
orbits).  The  cranial  elements  are  deeply  marked  with  pits  and  short,  shallow 
grooves.  On  the  left  mandible  these  pits  are  in  a  very  distinct  row,  the  operculo- 
mandibular  lateral  line. 


ij £?*'** 


Macrerpeton  deani  new  species. 

Type :  Specimen  No.  2934,  American  Museum  of  Natural  History. 

Horizon  and  locality:  Linton,  Ohio,  Coal  Measures.     (Plate  21,  figs.  1,2.) 

The  material  for  this  species  consists  of  the  posterior  half  of  the  left  mandible 
and  a  portion  of  the  right  antero-lateral  surface  of  the  skull,  both  incomplete.  The 
reasons  for  regarding  the  species 
as  distinct  are  the  large  size  of 
the  specimens  and  the  manner  of 
the  sculpture,  as  well  as  the 
shape  of  the  posterior  end  of  the 
mandible. 

The  present  species  is  the 
largest  amphibian  of  the  Linton, 
Ohio,  Coal  Measures,  exceeding 
in  skull  length  that  of  Macrer- 
peton huxleyi  by  twice.  The 
largest  skull  of  Macrerpeton  huxleyi  which  has  so  far  come  under  my  notice  is  120 
mm.  in  median  length.  There  are  3  skulls  of  this  species  known,  all  of  approxi- 
mately the  same  size.  The  skull  of  Macrerpeton  deani  must  have  reached  or  exceeded 
a  foot  in  median  length.  The  only  species  with  which  it  can  at  all  be  com- 
pared are  Eobaphetes  kansensis  Moodie  and  Baphetes  planiceps  Owen,  but  it  is 
clearly  distinct  from  all  other  genera  of  Linton  Amphibia.  It  is  possible  that  when 
better  known  Macrerpeton,  Eobaphetes,  Baphetes,  Erpetosaurus,  and  possibly  Dend- 
rerpeton  will  form  a  natural  group  of  early  labyrinthodont-like  Amphibia. 

The  mandible  is  similar  in  structure  to  that  of  the  labyrinthodonts,  with  the 
elements  marked  by  radiate  flutings.     I  can  detect  no  evidences  of  a  lateral-line 


Fig.  40. — Mandible  of  Macrerpeton  deani  new  species,    from  Linton, 
Ohio.     X  0.75. 


THE   TEMNOSPONDYLOUS   AMPHIBIA.  1 85 

canal,  such  as  is  clearly  marked  in  Macrerpeton  huxleyi  Cope  by  a  series  of  rounded 
pits,  occupying  the  usual  position  of  the  operculo-mandibular  lateral-line  canal 
The  teeth,  of  which  6  are  preserved,  are  minutely  striate,  with  smooth  apices.    They 
are  dissimilar  in  size,  showing  a  variation  of  2  or  3  mm.  in  length. 

The  sculpture  is  a  coarse  fluting,  with  no  indications  of  the  sharply  marked  pits 
of  Macrerpeton  huxleyi  Cope. 

The  fragment  of  a  skull  preserved  shows  characters  of  the  sculpture  which  are 
identical  with  those  of  the  mandible.  The  bones  are  so  crushed  that  it  is  impossible 
to  tell  the  limits  of  the  elements.  I  believe  a  portion  of  one  orbit  is  represented  on 
one  corner  of  the  block.  The  cranium  appears  to  have  been  broad,  and  the  fragment 
preserved,  which  is  only  about  one-sixth  of  the  skull,  is  larger  than  the  entire 
cranium  and  mandibles  of  Macrerpeton  huxleyi. 

The  specific  distinctness  of  the  form  can  not  be  doubted,  although  it  is  a  mat- 
ter of  regret  that  it  is  founded  on  so  small  a  portion  of  the  osteology  of  the  animal. 

The  species  is  proposed  in  honor  of  Dr.  Bashford  Dean,  to  whom  I  am  greatly 
indebted  for  many  kindnesses  during  the  past  5  years  in  connection  with  my  studies 
on  Carboniferous  Amphibia,  particularly  in  the  loan  of  the  entire  Newberry  col- 
lection of  Linton,  Ohio,  Amphibia. 

Measurements  of  the  Type  of  Macrerpeton  deani  Moodie. 

mm. 

Length  of  portion  preserved 115 

Greatest  width 50 

Ix'nglh  of  tooth 9 

Width  at  base 4 

Length  of  angular 95 

Diameter  of  angular 25 

Measurements  of  specimen  No.  8535  G,  American  Museum  of  Natural  History,  associated 
with  the  above  in  the  type  description: 

Length  of  preserved  portion 140 

Diameter  of  orbit 22 

Family  ANTHRACOSAURIDjE  Cope,  1875. 

Cope,  Bull.  U.  S.  Nat.  Mus.,  I,  p.  10,  1875. 

Lydekker,  R.,  1890,  Cat.  Fossil  Reptilia  and  Amphibia,  p.  157. 

Skull  usually  triangular  and  more  or  less  angulated,  with  the  cranial  sculpture 
well  marked,  the  occipital  condyles  ossified,  and  the  palatine  foramina  very  small 
and  placed  far  back;  dentine  of  the  teeth  more  or  less  complexly  plicated.  A 
ventral  armor  of  elongated  dermal  scutes,  and  probably  a  sclerotic  ring.  Bodies 
of  vertebras  fully  ossified  in  the  adult ;  intercentra  present  or  absent.  According 
to  Atthey's  figure  (11)  of  the  skull  of  the  type  genus,  the  palatine  bears  teeth 
which  are  situated  immediately  on  the  inner  side  of  the  maxilla,  as  in  Masto- 
donsaurus  (242).  In  the  typical  forms  there  is  no  postarticular  process  to  the 
mandible. 

The  North  American  species  of  this  family  are:  Eosaurus  acadianus  Marsh, 
Eobaphetes  kansensis  Moodie,  Dendrerpeton  acadianum  Owen,  Dendrerpeton  oweni 
Dawson,  Platystegos  loricatum  Dawson,  Baphetes  planiceps  Owen,  Baphetes  minor 
Dawson. 

There  is  but  little  assurance  that  any  of  these  species  belong  in  this  family. 
They  are  put  there  provisionally,  pending  future  discoveries.  Huxley  suggests  the 
relationship  of  Eosaurus  and  Anthracosaur.us  (Quart.  Jour.  Geol.  Soc,  xix,  1863, 
p.  65;  Scientific  Memoirs,  11,  p.  566). 


1 86  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

Genus  BAPHETES  Owen,  1854. 

Owen,  Quart.  Jour.  Geol.  Soc.  London,  x,  p.  207,  pi.  ix,  1854. 
DAWSON,  Air-breathers  of  the  Coal  Period,  pp.  10-16,  pi.  ii,  1863. 

Type :  Baphetes  planiceps  Owen. 

Known  only  from  an  incomplete  skull,  which  is  large,  broader  than  long;  squa- 
mosals prolonged  into  obtuse  horns.  Teeth  rather  large,  heterodont,  arranged  in  a 
single  row.    Orbits  placed  well  forward,  frontals  small,  surface  bones  sculptured. 

Baphetes  planiceps  Owen. 

Owen,  Quart.  Jour.  Geol.  Soc.  London,  x,  p.  207,  pi.  ix,  1854. 
Dawson,  Air-breathers  of  the  Coal  Period,  pp.  10-16,  pi.  ii,  1863. 

Type :  Specimen  in  the  British  Museum  of  Natural  History. 

Horizon  and  locality:  Near  Pictou,  Nova  Scotia  (Coal  Measures). 

The  parts  preserved  include  the  premaxillaries,  nasals,  and  portions  of  the 
frontal,  prefrontal,  and  maxillary  bones.  The  fossil  is  embedded  in  a  mass  of  Pictou 
Coal  from  Nova  Scotia  and  consists  of  the  anterior  extremity  of  the  cranium  (plate 
22,  fig.  6)  and  with  the  exterior  surface  of  the  bone  embedded  in  the  matrix,  and  its 
substance,  for  the  most  part,  reduced  to  a  thin  layer  by  abrasion  of  the  exposed 
inner  layer.  It  displays  accurately  the  contour  of  the  fore  part  of  the  upper  jaw, 
which  was  broad,  obtuse,  and  rounded. 

The  premaxillaries,  which  show  some  obscure  traces  of  a  symphysial  suture  at 
the  median  line,  anterior  to  the  nasal  or  naso-palatine  vacuities,  extend  outwards, 
on  each  side,  for  an  extent  of  2.5  lines  and  there  join  the  maxillaries.  Traces  of 
round  alveoli  for  teeth,  some  of  which  are  2  lines  in  diameter,  are  visible  on  the 
alveolar  border  of  the  premaxillaries.  The  alveolar  border  is  continued  by  the  max- 
illary bone  for  an  extent  of  4.5  inches  beyond  the  premaxillary  border,  and  this 
border  shows  still  more  distinct  traces  of  alveoli,  of  a  circular  form,  about  a  line  in 
diameter  and  rather  close  set  in  a  single  series.  The  fore  part  of  the  orbit  is  very 
unequivocally  displayed,  the  smooth  inner  or  under  surface  of  the  bone  forming 
that  part  being  entire;  and  this  shows  the  fore  part  of  the  orbit  to  be  formed,  partly 
by  the  maxillary,  partly  by  the  lacrimal  or  prefrontal  bone  in  close  sutural  union 
therewith,  a  structure  which  does  not  exist  in  any  recent  or  fossil  fish  with  a  dentig- 
erous  superior  maxillary  bone.  Where  the  substance  of  the  bone  has  been  detached 
so  far  as  to  expose  the  external  layer  in  contact  with  the  coal,  as,  e.g.,  on  the  frontal 
and  part  of  the  prefrontal  bones,  the  external  surface  of  those  bones  is  shown  to 
have  been  impressed  by  subhemispherical  or  elliptical  pits,  from  1  line  to  1.5  lines 
in  diameter,  and  with  intervals  of  half  of  that  extent.  This  coarsely  pitted  character 
agrees  with  that  presented  by  the  other  surface  of  the  similarly  broad  and  flat 
cranium  of  the  labyrinthodonts. 

From  the  characters  above  specified,  therefore,  I  conclude  that  this  fossil  is 
the  fore  part  of  the  skull  of  an  extinct  family  of  the  labyrinthodonts.  It  agrees  with 
them  in  the  number,  size,  and  disposition  of  teeth ;  in  the  proportions  and  mode  of 
connection  of  the  premaxillaries,  maxillaries,  nasals,  prefrontals,  and  frontals,  and  in 
the  resultant  peculiarly  broad  and  depressed  character  of  the  skull.    The  traces  of 


THE   TEMNOSPONDYLOUS   AMPHIBIA.  1 87 

the  nostrils  are  less  definite  and  satisfactory  than  the  remains  of  the  orbits,  but  the 
latter  appear  to  be  decisive  against  the  piscine  nature  of  the  fossil.  The  fossil  also 
presents  the  same  well-marked  external  sculpturing  as  in  the  labyrinthodonts;  and 
among  the  genera  that  have  been  established  in  that  family,  the  form  of  the  end  of 
the  muzzle,  or  upper  jaw,  in  the  Pictou  coal  specimen  best  accords  with  that  in  the 
Capitosaurus  and  Metopias  of  von  Meyer  and  Burmeister  (80). 

Measurements  of  the  Skull  of  Baphetes  planiceps  Owen. 
(Type  in  the  British  Museum  of  Natural  History,  London.) 

mm.  mm. 

Approximate  median  length  of  skull 136      Width  of  skull  across  orbits 97 

Width  of  skull  across  base  of  horns 150      Diameter  of  orbit 21 

Estimated  width  across  tips  of  horns 186      Diameter  of  large  tooth  alveolus 7 

Width  of  horn  at  base 31      Diameter  of  small  tooth  alveolus 2 

Estimated  length  of  horn  from  base 80 

Pictou  Coal,  near  Pictou,  Nova  Scotia,  Canada,  collected  by  Dr.  J.  William 
Dawson,  1850,  and  presented  by  him  to  the  Geological  Society  of  London. 

Baphetes  minor  Dawson. 
Dawson,  Canadian  Nat.  and  Jour.  Sci.,  n.  s.,  1870,  v,  pp.  98,  99. 

Type:  Specimen  in  the  Peter  Redpath  Museum,  McGill  University. 

Horizon  and  locality:  Coal  formation  of  Nova  Scotia. 

The  species  was  based  on  a  lower  jaw  of  an  amphibian,  of  which  a  cast  had 
occurred  in  the  coarse  sandstone  of  the  coal  formation  between  Ragged  Reef  and 
the  Joggins  Coal  Mine.  It  measured  6  inches  in  length;  its  surface  was  marked  on 
the  lower  and  posterior  part  with  a  network  of  ridges  inclosing  rounded  depressions. 
The  anterior  part  of  the  jaw  contained  about  16  teeth,  some  of  which  remained  in 
the  matrix.  These  were  stout,  conical  and  blunt,  with  large  pulp  cavities,  and 
about  32  longitudinal  striae,  corresponding  to  the  folds  of  the  dentine.  Dawson 
states  that  this  jaw  resembles  most  closely  those  of  Baphetes  and  Dendrerpeton,  but 
more  especially  the  former.  He  regarded  it  as  distinct  from  Baphetes  planiceps, 
and  proposed  for  it  the  name  Baphetes  minor. 

Eosaurus  acadianus  Marsh. 

Marsh,  Am.  Jour.  Sci.  (2),  xxxiv,  pp.  1-16,  pis.  i,  ii,  1862. 

Agassiz,  Am.  Jour.  Sci.,  xxxm,  p.  138,  1862. 

Marsh,  Quart.  Jour.  Geol.  Soc,  xix,  pp.  52-56. 

Hay,  Cat.  Fossil  Vertebrates  (Bull.  U.  S.  Geol.  Surv.  No.  179,  p.  421,  1902). 

Type:  Specimen  No.  1648,  Yale  University  Museum. 

Horizon  and  locality:  South  Joggins,  Nova  Scotia  (Coal  Measures). 

The  genus  and  species  are  founded  on  two  vertebral  bodies  of  the  stereospondy- 
lous  type  from  the  Coal  Measures  of  the  South  Joggins,  Nova  Scotia.  Marsh's 
description  (404)  is  as  follows: 

"The  general  form  of  the  vertebrae  is  cylindrical,  but  their  sides  are  compressed  ob- 
liquely, which  gives  to  the  contour  of  the  centra  a  subhexagonal  appearance.  They  are 
much  flattened  in  the  direction  of  the  anteroposterior  diameter,  which  has  to  the  transverse 


i88 


THE   COAL   MEASURES    AMPHIBIA    OF    NORTH    AMERICA. 


diameter  the  proportion  of  i  to  3.  Both  the  articular  terminal  facets  are  deeply  and  equally 
concave;  but  from  the  center  to  the  margin  the  surfaces  are  convex,  and  this  convexity  is 
greatest  near  the  center.  *  *  *  The  cavities  for  the  reception  of  the  intervertebral  matter 
begin  immediately  from  the  margin,  and  are  considerably  deeper  than  the  corresponding 
parts  of  the  Ichthyosaurus,  indicating  a  greater  degree  of  flexibility  in  the  vertebral  column. 
The  margins  of  the  vertebras  are  some.what  raised,  as  if  they  had  yielded  to  a  forcible  com- 
pression applied  longitudinally  ;  and  hence  the  lateral  surfaces  of  the  centers  are  concave  in 
an  anteroposterior  direction.  This  concavity  is  greater  in  the  upper  half  of  the  vertebra 
and  was  undoubtedly  more  marked  originally  than  at  present,  since  the  appearance  of  the 
margins  indicates  considerable  abrasion.  The  non-articular  surfaces  of  the  centra  are 
smooth  and  regular;  and  the  external  fibres  of  the  osseous  tissue  are  singularly  reticulated. 


(Mi) 

\v\  ♦*c 

F 

Fig.  41. — Nova  Scotian  Amphibia. 

A.  Oblique  lateral  view  of  vertebra?  of  Eosaurus  acadianus  Marsh,    a,  pits 

for  articulation  of  neuropophy ses ;  b,  rudimentary  transverse  process  on 
right  lateral  surface  of  centrum. 

B.  Oblique  view  of  vertebra?,    a,  pits  for  articulation  of  neuropophy  ses ;  6, 

rudimentary  process  on  lateral  surface. 

C.  Posterior  view  of  nearly  perfect  centrum. 

D.  Transverse  section  of  same  vertebra,  showing  deep  concavities  of  articular 

terminal  facets. 
E  and  F.  Microscopic  sections  near  surface,  showing  lacuna?  arranged  around 

an  Haversian  canal.    Magnified  200  diameters. 
(All  figures  after  Marsh.     X  0.75.) 


'  The  neuropophyses  are  not  anchylosed  to  the  centrum,  as  in  the  mammalia,  nor  con- 
nected to  it  by  sutures,  as  in  the  crocodiles ;  but  their  union  with  the  vertebra  is  indicated  by 
two  pits,  which  served  for  their  articulating  surfaces.  These  depressions  are  situated  on 
the  superior  surface  of  the  centrum  intermediate  between  the  anterior  and  posterior  mar- 
gins of  the  extremities.  They  are  circular  in  form  and  sink  directly  into  the  body  of  the  ver- 
tebra; instead  of  being  elongated  longitudinally  and  raised  by  ridges,  as  in  Ichthyosaurus. 
The  pits  are  about  a  line  in  depth,  and  in  the  more  perfect  of  the  fossils  are  not  in  their  origi- 
nal position.  The  floor  of  the  spinal  canal  is  narrow,  being  but  5  lines  in  breadth.  A  rudi- 
mentary transverse  process,  or  exogenous  tubercle,  is  sent  off  from  each  lateral  surface  of 
the  centrum,  at  points  equidistant  from  the  extremities  of  the  vertical  diameter.     Their 


THE    TEMNOSPONDYLOUS    AMPHIBIA.  1 89 

position  is  near  the  margin  of  the  anterior  articular  surface,  and  the  edges  of  these  para- 
pophyses  make  the  transverse  diameter  of  this  extremity  somewhat  greater  than  that  of 
the  corresponding  posterior  facet." 

Measurements  of  the  More  Perfect  Vertehra  of  Eosai  ms  acadianvs  (after  Marsh). 


mm. 


Transverse  diameter  of  centrum  on  anterior  surface 59 

Same  on  posterior  surface 57 

Same  including  the  parapophyses 63 

Vertical  diameter  of  anterior  surface P4 

Anteroposterior  diameter  on  superior  surface 21 

Same  on  inferior  surface ny 

Same  between  centers  of  articular  facets 2.5 

Length  of  pits  for  articulation  of  neural  arch 7 

Breadth  of  same 7 

Depth  of  same 2 

Distance  between  centers  of  same II 

Same  at  centers  of  parapophyses 36 

Collected  by  0.  C.  Marsh  at  the  South  Joggins,  Nova  Scotia. 

Marsh  regarded  (404)  the  vertebrae  as  representing  a  new  type  of  ichthyosaurian 
(2),  but  there  can  be  no  doubt  that  the  vertebras  belong  to  some  form  of  the  Amphibia, 
since  the  description  applies  equally  well  to  them.  In  this  connection  mention 
must  be  made  of  a  large  rib  from  the  Linton  beds  preserved  in  the  U.  S.  National 
Museum.  Only  the  proximal  third  of  the  rib  is  preserved,  but  it  represents  some 
large  form  of  the  Stegocephala.  The  rib  is  strongly  curved  backward,  is  heavy, 
and  has  an  incipient  tubercle.  A  cross-section  shows  that  a  longitudinal  groove 
occupies  the  median  line  on  the  exposed  surface  of  the  rib.  This  may,  however,  be 
due  to  compression  and  thus  indicate  that  the  rib  was  hollow.  The  rib  as  pre- 
served measures:  length,  102  mm.;  maximum  width,  22  mm.;  minimum  width,  14 
mm.    (Nos.  4490,  4489,  U.  S.  National  Museum.) 

Genus  E0BAPHETES  new  name. 

Type:  Eobaphetes  kansensis  MoOdie. 

The  new  name  is  proposed  to  replace  the  generic  term  Erpetosuchus  used  for  the 
species  E.  kansensis  described  by  the  writer  (Proc.  U.  S.  Nat.  Mus.,  39,  p.  491, 
191 1),  and  which  later  was  found  to  be  preoccupied  by  Newton  (Phil.  Trans.  Roy. 
Soc.  London,  185,  p.  573,  1894  b). 

The  genus  is  very  readily  distinguished  by  two  prominent  characters — the  short, 
uniform  dentition  and  the  presence  of  two  elongate,  oval,  internal  mandibular  fora- 
mina on  the  inner  side  of  the  jaw.  The  genus  may  be  further  distinguished  by  the 
great  depth  of  the  posterior  portion  of  the  jaw  and  the  slender  anterior  part,  as  well 
as  by  the  ornamentation,  which  is  typically  the  rough  tuberculated  labyrinthodont 
sculpture  on  the  anterior  end  of  the  mandible.  This  changes  gradually  to  longi- 
tudinal grooves  and  ridges  of  a  rather  small  size  on  the  posterior  portion,  a  very 
unusual  arrangement  for  a  labyrinthodont. 

These  characters  are  sustained  by  those  of  the  skull  fragment,  in  which  the  den- 
tition is  uniform  and  the  sculpture  very  similar  to  that  of  the  mandible.  The  ribs 
are  long,  curved,  and  solid,  as  in  other  labyrinthodonts. 


I90  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

The  internal  surface  of  the  mandible  shows  much  similarity  to  that  of  the  croco- 
diles and  alligators  of  the  present  day.  The  resemblance  is  not  due  to  homology  of 
structures,  but  must  be  regarded  as  a  parallel  development  of  similar  characters. 

Eobaphetes  kansensis  Moodie. 
Moodie,  Proc.  U.  S.  Nat.  Mus.,  39,  pp.  491-494,  figs.  1  to  3,  191 1  (Erpetosuchus). 

Type:  Specimen  No.  6699,  U.  S.  National  Museum. 

Horizon  and  locality:  Coal  Measures  of  Washington  County,  Kansas. 

The  species  is  represented  in  the  collections  of  the  U.  S.  National  Museum  by  a 
fragment  of  the  skull,  with  portions  of  two  ribs  (Cat.  No.  6699,  Vert.  Pal.  U.  S. 
Nat.  Mus.)  and  the  larger  part  of  the  left  ramus  of  the  mandible  (Cat.  No.  6680, 
Vert.  Pal.  U.  S.  Nat.  Mus.).  The  mandible  was  preserved  in  a  large  block  of  coal 
which  contained  the  impression  of  the  back  portion  of  the  mandible  from  which  the 
bone  had  been  weathered.  It  was  possible  to  remove  the  bone  and  make  a  plaster 
cast  of  the  impression.  This  shows  in  a  very  satisfactory  manner  all  of  the  char- 
acters of  the  external  surface. 

Skull. — Only  a  portion  of  the  left  maxilla,  with  14  teeth,  and  a  part  of  the  nasal 
are  preserved.  The  skull  seems  to  have  been  laterally  crushed  and  the  right  side 
of  the  skull  has  been  crushed  flat  under  the  left.  It  has  not  seemed  feasible  to 
remove  the  skull  from  the  matrix. 

The  teeth  are  uniform,  rather  short,  bluntly  conical,  curved  backward,  and 
coarsely  striate.  They  are  somewhat  crowded,  the  bases  being  separated  from  each 
other  by  only  a  fraction  of  a  millimeter. 

The  maxilla  and  portion  of  the  nasal  are  coarsely  sculptured  with  elongate  pits 
and  ridges.  A  portion  of  the  infraorbital  lateral-line  canal  is  preserved.  It  is  simply 
a  rounded  groove  with  three  short  branches.    It  lies  near  the  middle  of  the  maxilla. 

Mandible. — It  has  been  possible  to  study  both  sides  of  the  mandible.  The 
left  ramus  was  preserved  in  the  coal,  with  its  inner  face  exposed.  This  face  is  broken 
by  two  large  oval  openings,  the  internal  mandibular  foramina.  This  is  the  term 
used  by  Reynolds  for  the  openings  on  the  inner  surface  of  the  alligator  jaw.  So 
far  as  I  can  ascertain,  no  other  known  labyrinthodont  mandible  displays  this  char- 
acter in  such  a  marked  degree.  Dr.  Branson  has  figured  in  Anaschisma  browiri 
Branson  from  the  Triassic  (49)  of  Wyoming  the  inner  surface  of  the  left  ramus,  on 
which  there  are  likewise  two  openings  but  differently  situated.  A  similarity  be- 
tween the  two  mandibles  is  observed  in  that  the  suture  separating  the  prearticular 
and  angular  touches  the  posterior  edge  of  the  posterior  foramen. 

Several  of  the  sutures  are  well  preserved  and  they  have  been  indicated  in  the 
drawing  (fig.  42).  The  pillar  separating  the  two  foramina  is  cut  by  the  suture 
separating  the  angular  and  prearticular  very  much  as  in  Anaschisma,  with  the 
difference  that  in  the  latter  form  the  angular  and  prearticular  are  not  approximated. 
I  believe  I  detect  the  suture  as  represented  separating  the  anterior  end  of  the  angular 
from  the  dentary  and  splenial.  I  am  assured  of  the  portion  near  the  anterior  fora- 
men and  also  of  the  part  near  the  tip  of  the  ramus.  This  shows  the  angular  to  be  a 
very  elongate  element,  running  very  nearly  the  entire  length  of  the  mandible,  much 


THE    TEMNOSPONDYLOUS   AMPHIBIA. 


191 


as  in  Anaschisma  and  other  labyrinthodont  genera.  The  splenial  is  a  small,  slender 
element  located  farther  forward,  where  it  has  been  shoved  by  the  large-sized  internal 
mandibular  foramina.  The  prearticular  is  a  rather  long,  broad  element,  of  which 
only  a  portion  is  preserved.  I  am  not  sure  as  to  the  location  of  the  suture  for  the 
dentary,  unless  it  is  represented  by  the  line  bounding  the  roughened  area  near  the 
teeth.  If  this  is  true,  the  dentary  is  a  large  element,  since  it  extends  well  down  upon 
the  outer  side  of  the  jaw.  The  dentary  possesses  evidences  of  26  teeth,  a  few  of 
which  are  completely  preserved.  Most  of  them  are,  however,  represented  either 
by  bases  or  by  impressions  in  the  coal.  The  teeth  are  very  similar  to  those  of  the 
maxilla,  though  slightly  larger.  The  characters  given  for  the  maxillary  teeth  will 
suffice  for  those  of  the  dentary. 


*  1         \  - 

Fig.  42. 

A.  Outer  view  of  mandible  of  Eobaphetes  kansensis  Moodie,  from  the  Coal 

Measures  of  Washington  County,  Kansas.  Original  in  U.  S.  National 
Museum.     X  0.33.    Cat.  No.  6680.    o,  angular;  particular;  d,  dentary. 

B.  Portion  of  skull  of  Eobaphetes  kansensis  Moodie,  from  the  Coal  Meas- 

ures of  Washington  County,  Kansas.  Original  in  U.  S.  National 
Museum.  Cat.  No.  6699.  X  0.33.  Lateral-line  canal  represented  by 
heavy  broken  line,    n,  nasal;  m,  maxilla. 

C.  Inner  surface  of  mandible  of  Eobaphetes  kansensis  Moodie,  from  the 

Coal  Measures  of  Washington  County,  Kansas.  X  0.33.  a,  angular; 
p,  prearticular;  r,  articular;  s,  splenial. 

The  markings  of  the  inner  surface  are  as  indicated  in  the  drawing.  The  back 
portion  of  the  angular  shows  a  few  radiating  lines.  The  dentary  is  roughened  in  two 
portions:  one  near  the  teeth,  the  other  at  the  tip,  where  there  is  a  cartilaginous 
roughening  for  union  with  its  mate.  The  remainder  of  the  inner  surface  is  relatively 
smooth. 

The  outer  surface  shows  at  the  anterior  end  the  typical  labyrinthodont  sculptur- 
ing, which  becomes  slight  grooves  and  ridges  posteriorly.  I  detect  evidences  of  the 
operculo-mandibular  lateral-line  canal  throughout  the  entire  length  of  the  mandible. 
Its  location  is  indicated  by  the  heavy  broken  line.  The  suture  between  the  dentary 
and  angular  is  quite  clear.  The  suture  separating  the  dentary  and  splenial  joins 
the  angular  suture  about  midway  of  the  length  of  the  jaw. 

Measurements  of  Skull  Fragment  of  Eobaphetes  kansensis  Moodie. 

(Cat.  No.  6699,  U.  S.  Nat.  Mus.) 

mm.  rom. 

Total  length  of  portion  preserved 109      Length  of  tooth 10 

Maximum  width  of  maxilla 45      Width  of  tooth  at  base 4 

1  hickness  of  maxilla 7 


19-2  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 

Measurements  of  Left  Ramus  of  Mandible  of  Eobaphetes  kansensis  Moodie. 

(Cat.  No.  6680,  U.  S.  Nat.  Mus.) 

mm.  mm. 

Total  length  of  jaw,  as  preserved 305  Length  of  anterior  internal  mandibular  foramen. ...  56 

Greatest  width 79      Greatest  width 15 

Least  width 24      Least  width 7 

Length  of  angular 132  Length  of  posterior  internal  mandibular  foramen.  .  .  77 

Width  of  angular 45      Greatest  width 28 

Length  of  largest  tooth 10      Least  width 14 

Width  of  largest  tooth  at  base 6      Length  of  bridge 16 

Length  of  most  posterior  tooth 6      Width  of  bridge 8 

Width  of  most  posterior  tooth  at  base 4 

Ribs. — There  are  portions  of  two  dorsal  ribs  preserved  on  the  block  of  coal  with 
the  skull.  These  show  characters  very  similar  to  those  exhibited  by  the  rib  ascribed 
to  Macrerpeton  huxleyi  Cope,  and  also  those  of  Metoposaurus  diagnosticus  von  Meyer 
(242)  and  Anaschisma.  The  ribs  are  solid,  heavy,  curved,  and  have  a  longitudinal 
groove  on  the  middle  of  each  side.  The  heads  of  the  ribs  in  the  present  specimen  are 
obscured  and  nothing  can  be  said  of  them  except  that  they  appear  to  be  large. 

Measurements  of  Ribs  of  Eobaphetes  kansensis  Moodie. 

mm. 

Length  of  preserved  portion 130 

Width  at  distal  end 18 

Thickness  of  rib 5 

Genus  DENDRERPETON  Owen,  1853. 

Owen,  Quart.  Jour.  Geol.  Soc.  London,  ix,  p.  66,  1853. 

Type:  Dendrerpeton  acadianum  Owen. 

The  genus  is  characterized  by  Dawson  as  follows  (Phil.  Trans.,  1882,  p.  635): 

"  Lizard-like,  with  anterior  and  posterior  extremities  nearly  equal;  the  skull  somewhat 
elongate  with  small  orbits,  and  the  nostrils  placed  at  the  front.  The  cranial  bones  sculp- 
tured. The  teeth  plicated  at  the  base,  more  especially  on  their  inner  sides.  A  series  of  large 
teeth  on  the  palate.  The  body  was  covered  above  with  imbricated  horny  scales  and  had 
lappets  or  pendants  at  the  sides.  The  abdomen  was  protected  by  thin  bony  scales  semi- 
elliptical  or  oat-shaped  in  form,  and  arranged  in  a  chevron  pattern.  There  was  probably 
also  a  thoracic  plate.    Two  species:  D.  acadianum  and  D.  oweni. 

"Type:  D.  acadianum  Owen." 

Dendrerpeton  acadianum  Owen. 

Owen,  Quart.  Jour.  Geol.  Soc.  London,  ix,  p.  66,  1853. 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  17,  1863. 

Dawson,  Acadian  Geology,  3d  ed.,  p.  362. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  pt.  11,  p.  642,  pi.  40,  figs.  46  to  51;  pi.  44,  figs.  129  to  137.     1882. 

Type:  Specimens  Nos.  434-438,  British  Museum  of  Natural  History. 

Horizon  and  locality:  Coal  formation  at  South  Joggins,  Nova  Scotia.   (Plate  6.) 

This  species  has  been  fully  described  and  figured  by  Dawson  (Air-breathers  of 

the  Coal  Period,  pp.  17-30,  pi.  in,  figs.  1  to  30,  1863),  who  gives  a  detailed  account 

of  the  discovery  of  the  material  of  this  species  by  himself  and  Sir  Charles  Lyell. 

He  says,  in  part : 

"In  form,  Dendrerpeton  acadianum  was  probably  lizard-like;  with  a  broad  flat  head, 
short,  stout  limbs  and  an  elongated  tail;  and  having  its  skin,  and  more  particularly  that  of  the 
belly,  protected  by  small  bony  plates  closely  overlapping  each  other.  It  may  have  attained 
the  length  of  2  feet.    The  form  of  the  head  is  not  unlike  that  of  Baphetes,  but  longer  in  pro- 


THE   TEMNOSPONDYLOUS   AMPHIBIA.  I93 

portion ;  and  much  resembles  that  of  the  labyrinthodont  reptiles  of  the  Trias.  The  bones 
of  the  skull  are  sculptured  as  in  Baphetes,  but  in  a  smaller  pattern.  The  nostrils  are  small, 
and  near  the  muzzle ;  the  orbits  are  circular,  and  separated  by  a  space  of  more  than  their 
own  diameter.  In  the  upper  jaw  there  is  a  series  of  conical  teeth  on  the  maxillary  and  inter- 
maxillary bones.  Those  on  the  intermaxillaries  are  much  larger  than  the  others,  and  have 
the  aspect  of  canines  or  tusks.  Within  this  outer  series  of  teeth,  but  implanted  apparently 
in  the  same  bones,  there  is  as  in  Archegosaurus  a  second  series  of  teeth,  closely  placed,  or 
with  intervals  equal  to  the  diameter  of  one  tooth.  These  inner  teeth  are  longer  than  the 
others,  implanted  in  shallow  sockets,  to  which  they  are  anchylosed,  and  have  the  dentine 
plicated,  except  toward  the  point.  A  third  group  of  teeth,  blunt  at  the  points,  largely  hol- 
low in  the  interior,  and  with  the  dentine  quite  simple,  appears  in  detached  bones,  which 
may  represent  the  vomer.  Only  a  part  of  this  formidable  armature  of  the  teeth  appears  in 
the  skull,  as  the  bones  of  the  roof  of  the  mouth  have  been  removed,  adhering  to  the  opposite 
side  of  the  matrix ;  but  the  fact  of  the  occurrence  of  two  sets  of  teeth  was  ascertained  by 
Professor  Wyman,  from  the  original  specimens,  and  is  manifest  in  the  fragment  *  *  * 
while  the  other  teeth,  supposed  to  be  vomerine,  appear  in  fragments  which  must,  from  their 
size  and  collocation,  have  belonged  to  Dendrerpeton.  It  will  be  observed  that  all  of  these 
teeth  are  anchylosed  to  the  bone ;  and  that  those  of  the  vomer  are  thinly  walled  and  simple, 
the  outer  series  on  the  maxillaries  and  intermaxillaries  simple  and  flattened,  while  the  inner 
series  of  teeth  are  conical  and  plicated.  In  the  lower  jaw  there  was  a  uniform  series  of 
conical  teeth,  not  perceptibly  enlarged  toward  the  front;  at  least  this  is  the  case  in  the  only 
specimen  at  present  in  my  collection;  which  is,  however,  merely  an  imperfect  cast  in 
hard  sandstone. 

' '  The  scapular  and  sternal  bones  seem  to  have  been  well  developed  and  strong,  but  only 
portions  of  them  are  known.  The  fore  limb  of  the  adult  animal,  including  the  toes,  must 
have  been  4  or  5  inches  in  length,  and  is  of  massive  proportions.  The  bones  are  hollow,  and 
in  the  case  of  the  phalanges  the  bony  walls  were  thin,  so  that  they  are  often  found  crushed 
flat.  The  humerus,  however,  was  a  strong  bone,  with  thick  walls  and  a  cancellated  structure 
toward  its  extremities;  still,  even  these  have  sometimes  yielded  to  the  great  pressure  to 
which  they  have  been  subjected.  The  cavity  of  the  interior  of  the  limb-bones  is  usually 
filled  with  calc-spar  stained  with  organic  matter,  but  showing  no  structure;  and  the  inner 
side  of  the  bony  wall  is  smooth,  without  any  indications  of  cartilaginous  matter  lining  it. 

' '  The  vertebrae,  in  the  external  aspect  of  their  bodies,  remind  one  of  those  of  fishes, 
expanding  toward  the  extremities,  and  being  deeply  hollowed  by  conical  cavities,  which 
appear  even  to  meet  in  the  center.  There  is,  however,  a  large  and  flattened  neural  spine. 
The  vertebrae  are  usually  much  crushed,  and  it  is  almost  impossible  to  disengage  them  from 
the  stone.  *  *  *  in  its  long  neural  and  hasmal  spines,  reminds  us  of  the  caudal  vertebrae  of 
those  batrachians  and  reptiles  which  have  tails  flattened  for  swimming,  and  probably  indi- 
cates that  this  was  the  case  with  Dendrerpeton.  The  ribs  are  long  and  curved,  with  an 
expanded  head,  near  to  which  they  are  solid,  but  become  hollow  toward  the  middle ;  and  the 
distal  extremities  are  flattened  and  thin-walled.  The  posterior  seems  to  have  been  not 
larger  than  the  anterior,  perhaps  smaller.  The  tibia  is  much  flattened  at  the  extremity,  as 
in  some  labyrinthodonts,  and  the  foot  must  have  been  broad,  and  probably  suited  for  swim- 
ming or  walking  on  soft  mud,  or  both.  That  the  hind  limb  was  adapted  for  walking  is  shown 
not  merely  by  the  form  of  the  bones,  but  also  by  that  of  the  pelvis,  the  best  preserved  speci- 
men of  which  I  have  represented  (208,  pi.  in,  fig.  28). 

"The  external  scales  are  thin,  oblique-rhomboidal  or  elongated-oval,  marked  with 
slight  concentric  lines,  but  otherwise  smooth,  and  having  a  thickened  ridge  or  margin;  in 
which  they  resemble  those  of  Archegosaurus,  and  also  those  of  Pholidogasier .  *  *  *  The 
microscopic  structure  of  the  scales  is  quite  similar  to  that  of  the  other  bones,  and  different 
from  that  of  the  scales  of  ganoid  fishes  *  *  *  ." 


194  THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 

Dendrerpeton  oweni  Dawson. 

Dawson,  Quart.  Jour.  Geol.  Soc.  London,  xvm,  p.  460. 

Dawson,  Air-breathers  of  the  Coal  Period,  p.  32,  1863. 

Dawson,  Acadian  "Geology,  3d  ed.,  p.  368. 

Dawson,  Phil.  Trans.  Roy.  Soc.  London,  pt.  11,  p.  643,  pi.  44,  figs.  131,  138,  139,  1882. 

A  smaller  species  than  the  preceding.  The  form  (plate  13)  is  fully  described  by 
Dawson  (Air-breathers  of  the  Coal  Period,  p.  32,  pi.  iv,  1863)  as  follows: 

"Among  the  reptilian  remains  found  in  erect  trees  at  the  South  Joggins,  there  have 
occurred  several  portions  of  skeletons  which,  from  their  sculptural  cranial  bones,  plicated 
teeth,  and  the  forms  of  their  scales  and  limb-bones,  I  have  referred  to  the  genus  Dendrer- 
peton, but  to  individuals  of  much  smaller  size  than  the  full-grown  specimens  of  D.  acad- 
ianum.  It  did  not  occur  to  me  to  suppose  that  these  were  specifically  distinct  from  the 
larger  individuals,  until  I  observed  that  bones  of  this  kind,  contained  in  the  collections  sent 
by  me  to  the  Geological  Society,  or  represented  in  the  figures  drawn  to  illustrate  one  of  my 
papers,  were  referred  by  Professor  Owen,  in  his  notes  on  these  specimens  and  figures,  in  the 
Journal  of  the  Geological  Society,  to  the  genus  Hylonomus;  which  is  quite  distinct  from 
Dendrerpeton,  as  will  be  explained  in  sequel. 

' '  I  was  thus  induced  to  reexamine  all  the  specimens  in  my  collection  and  the  result  has 
been  to  establish  a  strong  probability  that  there  is  in  reality  a  second  species  of  Dendrer- 
peton, smaller  than  D.  acadianum,  and  differing  from  it  in  several  points.  This  species  I 
propose  to  name  D.  oweni.  It  differs  from  D.  acadianum  in  the  following  particulars :  (1) 
its  much  smaller  size;  (2)  its  long  and  hooked  teeth  (it  will  be  seen  that  these  teeth  differ 
very  markedly  in  their  proportions  and  form  from  those  of  the  larger  species)  ;  (3)  the  greater 
plication  of  the  ivory  in  the  intermaxillary  teeth  (in  D.  acadianum  these  teeth  are,  on  the 
outside,  simple  almost  to  the  base,  and  plicated  on  the  inner  side,  while  in  this  species  they 
are  plicated  all  around  like  the  inner  maxillary  teeth) ;  (4)  the  form  of  the  skull,  which  has 
the  orbits  larger  in  proportion,  and  is  also  shorter  and  broader.  On  the  other  hand,  when 
we  have  described  the  species  Hylonomus,  it  will  be  seen  that  this  animal,  except  in  size, 
differs  from  them  quite  as  widely  as  does  D.  acadianum. 

' '  The  distinctness  of  D.  oweni  is  further  confirmed  by  the  fact  that  I  possess  small 
jaw-bones  of  Dendrerpeton,-  about  the  size  of  those  of  this  species,  but  having  the  teeth 
similar  in  form  to  those  of  the  larger  species ;  these  I  suppose  to  have  belonged  to  young 
individuals. 

"On  examining  the  figures  (208,  pi.  iv)  it  will  be  seen  that  the  bones  of  the  skull  were 
corrugated  as  in  the  large  Dendrerpeton,  but  with  a  smaller  pattern.  The  forms  of  the  jaw- 
bones also,  and  of  the  vertebras,  ribs,  scapular  bone,  bones  of  the  limbs,  and  bony  scales,  are 
very  similar,  and  indicate  that  in  general  form  this  creature  was  not  far  removed  from  its 
larger  relative.  The  bones  of  the  foot  especially  deserve  attention.  This  is  the  most  per- 
fect foot  of  Dendrerpeton  hitherto  found;  and  I  have  enlarged  it  in  the  figure  (208)  in  order 
more  distinctly  to  show  its  parts.  It  presents  three  long  toes  with  traces  of  a  smaller  one  at 
each  side,  so  that  there  were  probably  five  in  all.  If  these  toes  be  compared  with  the  foot- 
prints on  the  slab  discovered  by  Dr.  Harding,  it  will  be  seen  that  they  very  closely  corre- 
spond, though  the  toes  of  the  present  species  are  much  smaller.  The  footprints  are  precisely 
those  which  we  may  suppose  an  animal  of  the  size  of  Dendrerpeton  acadianum  would  have 
made  if,  as  the  bones  found  render  in  every  way  probable,  this  larger  species  had  a  foot 
similar  to  that  of  D.  oweni.  I  suppose,  for  this  reason,  that  these  footprints  are  really  those 
of  Dendrerpeton  acadianum  and  that  this  species  continued  to  exist  from  the  time  of  the 
lower  Coal  Measures  to  the  period  when  those  higher  beds  of  the  series,  in  which  its  bones 
are  found  at  the  Joggins,  were  deposited. 

The  present  species  must  have  lived  in  the  same  places  with  its  larger  relative,  but 
may  have  differed  somewhat  in  its  habits.    Its  longer  and  sharper  teeth  may  have  been 


THE   TEMNOSPONDYLOUS   AMPHIBIA.  195 

better  suited  for  devouring  worms,  larvae,  or  soft-skinned  fishes,  while  those  of  the  larger 
Dendrerpeton  were  better  adapted  to  deal  with  the  mailed  ganoids  of  the  period,  or  with 
those  smaller  reptiles  which  were  more  or  less  protected  with  bony  or  horny  scales. 

REMAINS  OF  SKIN  AND  HORNY  SCALES. 

' '  In  one  of  my  earliest  explorations  of  the  reptile-bearing  stumps  of  the  Joggins,  I 
observed  on  some  of  the  surfaces  patches  of  a  shining  black  substance,  which  on  minute 
examination  proved  to  be  the  remains  of  cuticle,  with  horny  scales  and  other  appendages. 
The  fragments  were  preserved;  but  I  found  it  impossible  to  determine  with  certainty  to 
which  of  the  species  whose  bones  occur  with  them  they  belonged,  or  even  to  ascertain  the 
precise  relations  of  the  several  fragments  to  each  other.  I  therefore  merely  mentioned  them 
in  general  terms,  and  stated  my  belief  that  they  may  have  belonged  to  the  species  of  Hylono- 
mus *  More  recently  other  specimens  have  been  obtained,  and  I  have  undertaken  the 
detailed  examination  of  the  whole.  I  shall  now  endeavor  to  describe  the  principal  or  most 
continuous  fragments,  and  afterward  to  consider  the  probabilities  of  their  having  belonged 
to  certain  of  the  reptiles  entombed  with  them.  I  do  this  here,  rather  than  under  the  titles 
of  these  several  animals,  on  account  of  the  uncertainty  which  still  rests  on  the  assignment 
of  certain  portions  of  this  cuticle  to  the  species  in  question,  and  which  renders  it  more  con- 
venient to  consider  these  peculiar  remains  in  one  place  and  to  compare  the  different  portions 
with  each  other. 

"(i)  One  of  my  specimens  is  a  flattened  portion  of  cuticle  2.25  inches  in  length.  The 
greater  part  of  the  surface  is  smooth  and  shining  to  the  naked  eye,  and  under  the  micro- 
scope shows  only  a  minute  granulation.  A  limited  portion  of  the  upper  and,  I  suppose, 
anterior  part  is  covered  with  imbricated  scales,  which  must  have  been  membranous  or  horny 
and  generally  have  a  small  spot  or  pore  near  the  outer  margin,  some  having  in  addition 
smaller  scales  or  points  on  their  surfaces.  In  contact  with  the  upper  part  of  this  specimen 
there  were  many  fragments  of  the  skull  of  Dendrerpeton  oweni. 

"(2)  Another  portion  of  the  cuticle,  similarly  marked,  appears  to  preserve  the  form  of 
the  posterior  part  of  the  body  and  tail  of  the  animal,  and  also  a  mark  representing  the  point 
of  attachment  of  the  hind  leg;  near  to  which,  and  along  the  dorsal  ridge,  is  a  portion  of  the 
skin  covered  with  much  smaller  scales.  This  was  found  in  close  proximity  to  a  mass  of 
bones  of  Dendrerpeton  oweni,  mingled  with  some  of  Hylonomus  lyelli. 

"(3)  A  third  and  still  larger  surface  of  integument  with  similar  markings  has  upon 
it  a  number  of  vertebras  and  detached  bones  of  the  small  reptile  Hylonomus  wymani, 
to  be  described  in  the  sequel;  for  which  species,  however,  it  would  be  much  too  large  a 
covering. 

"(4)  Another  well-preserved  fragment,  less  than  2  inches  in  length,  exhibits  very  dif- 
erent  markings.  It  is  nearly  covered  with  very  small  imbricated  scales,  thicker  than  those 
on  the  specimens  previously  described.  On  either  side  of  what  seems  to  have  been  the  mid- 
dle line  of  the  back,  there  is  a  series  of  pointed  flat  horny  processes,  which  probably  formed  a 
double  spinous  crest.  Without  these  there  are  tufts  of  strong  bristles,  and  exteriorly  to 
these  last  are  rows  of  flat,  thick,  horny  plates,  transversely  wrinkled.  Near  to  these  was  a 
row  of  conical  truncated  tubercles.  Sections  of  these  appendages  show  them  to  have  been 
horny  and  attached  to  the  cuticle.    None  of  them  have  bony  structure. 

"(5)  Near  this  last  portion  of  cuticle,  and  possibly  belonging  to  it,  are  pointed  and 
probably  membranous  appendages,  marked  on  each  side  with  rows  of  scales  not  overlapping 
and  each  with  a  pore  in  its  center.  The  manner  in  which  these  appendages  are  bent  and 
wrinkled  shows  that  they  must  have  been  soft,  except  at  the  tips,  which  seem  to  have  been 
hard  and  horny,  and  they  are  arranged  in  series,  as  if  originally  placed  along  the  sides  of  the 

*  Journal  of  Geological  Society,  vol.  xvi. 


I96  THE   COAL   MEASURES   AMPHIBIA   OF    NORTH    AMERICA. 

neck  or  abdomen,  or  both.  The  use  of  these  appendages  is  not  easy  to  conjecture.  They 
remind  us  of  the  gular  pouches  of  iguana,  and  of  the  lateral  expansions  of  some  geckos  and 
of  the  Draco  volans.  Possibly  they  formed  lateral  parachutes,  aiding  the  animal  in  moving 
over  soft  mud,  or  perhaps  in  leaping  and  swimming. 

"(6)  Some  other  fragments  appear  to  have  belonged  to  a  different  species  from  either 
of  the  foregoing.  The  best-preserved  specimen,  which  is  about  1  inch  in  length  and  half 
an  inch  in  breadth,  is  covered  with  very  small  imbricated  scales.  It  is  crossed  by  6  or  7 
obscure  ridges,  which  both  at  the  bottom  and  along  a  mesial  line  project  into  points 
covered  with  larger  scales.  A  row  of  large  scales  with  round  pores  connects  these  along 
the  lower  side.  If,  as  seems  probable,  this  fragment  belonged  to  the  side  of  the  trunk  or 
tail,  it  would  perhaps  indicate  a  division  of  the  subcutaneous  muscles  into  an  upper  and 
lower  band,  as  in  the  newts.  A  separate  fragment  with  transverse  horny  ridges  and 
another  with  a  longer  lobe,  similar  in  structure  to  those  above  mentioned,  may  perhaps 
be  referred  to  the  same  animal.  A  larger  patch  of  skin  presents  similar  imbricated  scales, 
but  without  a  mesial  line,  and  with  an  edging  of  larger  scales. 

' '  Six  species  of  reptiles  have  left  their  bones  in  the  repositories  containing  these 
remnants  of  cuticle.  Of  these,  Dendrerpeton  acadianum,  was  an  animal  of  too  great  size 
to  have  been  clothed  with  integument  of  this  character  and  of  such  dimensions.  Hylono- 
mus aciedentatus  and  Hylerpeton  dawsoni  are  each  represented  by  only  a  single  specimen, 
and  these  did  not  occur  in  proximity  to  any  of  the  portions  of  cuticle,  except  that  the 
appendages  were  found  near  a  specimen  of  the  former.  Of  the  three  remaining  species, 
Dendrerpeton  oweni,  from  its  size,  the  number  of  specimens  found,  and  the  juxtaposition 
of  their  bones  to  the  fragments  of  cuticle,  appears  to  have  the  best  claim  to  the  integu- 
ment included  under  Nos.  1,  2,  and  3;  and  in  this  case,  while  the  creature  had  its  throat, 
and  perhaps  its  abdomen,  armed  with  bony  scales,  its  upper  parts  and  tail,  as  well  as  its 
limbs,  had  a  uniform  covering  of  small,  thin  imbricated  horny  scales,  in  the  manner  of 
many  modern  reptiles. 

"  If  the  remaining  portions  of  integument,  Nos.  4  and  5,  as  would  seem  likely,  belonged 
to  two  species,  both  of  smaller  dimensions,  there  would  seem  little  reason  to  doubt  that 
these  were  Hylonomus  lyelli  and  H.  wymani.  In  this  case,  both  of  these  species  must  have 
possessed  a  highly  ornate  covering  of  horny  scales  and  appendages,  comparable  with  that 
of  many  of  the  modern  lizards,  while  there  seems  good  reason  to  believe,  as  stated  in  a  pre- 
vious paper,  that  they  were  in  part  protected  by  bony  scales  somewhat  like  those  of  Den- 
drerpeton. These  points,  however,  we  shall  consider  more  in  detail  under  the  sections 
which  refer  to  the  species  of  Hylonomus. 

Before  leaving  these  curious  specimens  of  ancient  skin,  the  most  ancient  I  suppose 
known  to  exist,  it  is  of  interest  to  observe  that  the  thicker  portions,  when  broken  across, 
have  the  aspect  of  jet,  or  of  pure  shining  coal,  and  thin  slices,  under  the  microscope,  have 
the  same  rich  brown  colour  with  that  material,  though  rather  more  translucent.  When 
burned,  fragments  of  the  substance  give  a  strong  flame,  and  a  bituminous  and  ammoniacal 
odour.  We  have  thus  an  example  of  the  production  of  coal  from  animal  membrane,  no 
doubt  gelatinous  and  horny  in  the  first  instance,  but  which  has  proved  itself  capable  of 
the  same  chemical  changes  that  have  been  experienced  by  the  vegetable  matter  buried 
with  it.  In  order  that  this  substance  should  be  preserved  in  this  way,  it  would  be  neces- 
sary that  it  should  either  be  kept  dry  and  hard,  or  that  it  should  be  immediately  buried 
in  matter  impervious  to  air  and  kept  moist.  The  latter  conditions  are  the  more  probable. 
The  preservative  qualities  of  the  peaty  vegetable  matter  imbedded  with  it  must  be  con- 
sidered ;  and  it  is  possible  that  these  hollow  stumps,  partly  filled  with  fragments  of  Sigillaria 
bark,  may  have  formed  natural  tan-pits,  in  which  animal  membranes  would  be  preserved 
in  a  manner  impossible  in  ordinary  sediments.  If  this  were  the  case,  we  may  yet  find  an 
entire  reptile,  preserved  as  a  flattened  mummy,  in  one  of  these  strange  repositories." 


THE   TEMNOSPONDYLOUS   AMPHIBIA.  197 

Genus  PLATYSTEGOS  Dawson,  189S. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  1895,  xn,  p.  77  (sec.  iv). 

Type:  Platystegos  loricatum  Dawson. 
Dawson's  description  is  as  follows : 

"Head  broad  and  short,  orbits  very  large,  cranial  bones  deeply  sculptured;  teeth 
strongly  plicated  and  curved,  with  sharp  edges  at  apices,  especially  the  inner  palatal  teeth, 
which  are  very  large.  Many  minute  teeth  on  the  vomerine  bones;  vertebrae  ossified,  bicon- 
cave ;  limb  bones  imperfectly  ossified,  short ;  lower  surface  protected  with  a  thoracic  plate 
and  thick,  densely  imbricated  bony  scales  in  transverse  rows;  body  above  with  thin,  round 
scales,  concentrically  marked." 

Platystegos  loricatum  Dawson. 
Dawson,  Proc.  and  Trans.  Roy.  Soc.  Canada,  xn,  p.  77,  1895. 

Dawson  describes  the  species: 

"  Head  about  8  cm.  long;  when  flattened  9  cm.  broad  across  parietal  foramen;  squa- 
mosal and  temporal  bones  projecting  backwards  in  points  much  behind  the  condyles; 
parietal  foramen  small,  orbits  large;  length  of  longest  tooth  seen  7  mm.;  cranial  bones 
closely  and  deeply  pitted;  humerus  with  very  thin  bony  walls,  cartilaginous  within,  3.5  cm. 
long." 

Erect  tree,  coal  formation,  at  the  South  Joggins;  collected  by  P.  W.  McNaugh- 
ton.  Type  in  Peter  Redpath  Museum  at  McGill  University.  Dawson  regarded 
the  form  as  of  uncertain  relationship. 


CHAPTER  XXV. 

THE  STEREOSPONDYLOUS  AMPHIBIA  FROM  THE  COAL  MEASURES 

OF  NORTH  AMERICA. 

DEFINITION  OF  THE  ORDER  STEREOSPONDYLIA  ZITTEL,  1887. 

Zittel,  Handbuch  der  Paleontologie,  Bd.  m,  Abth.  I,  p.  397,  1887. 

Large  terrestrial  vertebrates;  largest  of  the  class.  Skull  equal  to  one-fourth 
or  one-third  of  the  entire  body  in  at  least  one  species,  Metoposaurus  diagnosticus 
von  Meyer  (242).  Lateral-line  canals  always  present  (458)  on  the  skulls  as  deeply 
impressed  grooves  which,  in  life,  were  possibly  roofed  over  by  a  cartilaginous  or 
other  connective-tissue  membrane.  The  sensory  organs  undoubtedly  being  sup- 
plied by  the  superficial  ophthalmic  branch  of  the  trigeminal  nerve,  branches  of 
which  pierced  the  cranial  elements  near  the  grooves,  no  evidence  of  such  openings 
in  the  bottoms  of  the  grooves ;  the  condition  probably  being  analogous  to  Hydrola- 
gus  colei  and  other  chimaeroids.  Vertebras  stereospondylous,  with  well-developed 
neural  arches  from  which  projected  the  well-developed  zygapophyses,  sometimes 
slightly  amphiccelous  and  pierced  for  the  notochord,  such  forms  being  uncertainly 
placed  in  the  group.  Tail  unknown,  possibly  short.  Limbs  and  girdles  well 
developed  (243),  phalangeal  formula  unknown;  carpus  osseous  and  tarsus  un- 
known. Pectoral  girdle  composed  of  osseous  scapulae,  clavicles,  interclavicle, 
coracoid  (?);  clavicles  and  interclavicle  ventrally  sculptured.  Pelvic  girdle  com- 
posed of  osseous  pubis,  ischium,  and  ilium  (242),  the  pubis  a  small  plate,  in  life 
largely  cartilaginous,  the  three  uniting  by  cartilaginous  union  to  form  the  acetabu- 
lum.   Ventral  armature  unknown,  possibly  wanting. 

Range :  Pennsylvanian  to  Upper  Triassic. 

Distribution:  North  America,  Europe,  Asia,  Africa,  and  Australia. 

Family  MASTODONSAURIDjE  Huxley,  1863. 

Huxley,  Quart.  Jour.  Geol.  See,  xix,  p.  65,  1863. 

Lydekker,  R.,  Cat.  Fossil  Reptilia  Amphibia,  pt.  iv,  p.  141,  1890. 

Skull  triangular,  and  more  or  less  elongated,  with  the  cranial  bones  very 
strongly  sculptured,  the  occipital  condyles  ossified  (49),  and  large  palatal  vacuities; 
dentine  of  teeth  with  very  complex  plications  (502) ;  no  bony  rings  (242)  in  scle- 
rotic; and  no  ventral  scutes.  Bodies  of  vertebrae  (49)  fully  ossified  in  the  adult. 
There  are  large  palato-vomerine  tusks  on  the  inner  side  of  the  maxillary  teeth; 
and  the  palatines  run  parallel  to  the  maxilla.  The  mandible  has  a  large  post- 
articular  process;  and  there  is  a  small  inner  series  of  mandibular  teeth.  In  the 
type  genus  the  pubes  are  separate  from  the  ischia,  and  do  not  enter  into  the 
formation  of  the  acetabulum;  and  the  sacral  ribs  form  kidney-shaped  disks  (393). 

Represented  in  North  America  by  a  single  tooth  from  the  Carboniferous  of 
Kansas.  Described  by  Williston  as  Mastodonsaurus  sp.  (Kans.  Univ.  Quart.,  vi,  pp. 
209-210,  pi.  xxi).  Represented  in  the  Triassic  of  Wyoming  by  Anaschisma  browni, 
Branson  (49). 

Mastodonsaurus  sp.  indet. 
Williston,  Kans.  Univ.  Quart.,  vi,  p.  209,  1897. 

The  specimen  preserved  comprises  the  entire  crown  of  a  single  vomerine  (?) 

tooth,  38  mm.  in  length  by  14  mm.  in  diameter  at  base  (pi.  2 1 ,  fig.  6) .    The  immediate 

tip  had  been  partly  worn  away  in  life,  but  was  acuminate.    It  is  composed  of  a  dense 

198 


THE    STEREOSPONDYLOUS   AMPHIBIA.  199 

blackish  material,  with  the  exterior  smooth,  shining  black.  It  has  about  20  narrow 
flutings,  nearly  straight,  running  from  the  base  to  the  tip,  separating  shallow  grooves. 
A  transverse  section  of  the  base  shows  a  narrow  pulp-cavity  not  more  than  5  mm. 
in  diameter  which  extends  in  about  the  same  proportional  width  to  beyond  the 
middle  of  the  tooth,  and  in  all  probability  to  near  the  apex.  The  cross-section  of 
the  tooth  throughout  is  nearly  or  quite  circular. 

A  hemisection  of  the  tooth  was  made  near  the  middle,  showing  a  structure  most 
remarkably  like  that  of  Mastodonsaurus;  so  nearly  alike,  in  fact,  that  there  is  no  dif- 
ference from  the  large  figure  given  by  Owen  of  a  section  of  Mastodonsaurus  (502). 

The  discovery  of  this  tooth  in  the  Kansas  Coal  Measures  is  of  great  interest, 
proving,  as  it  does,  the  presence  of  true  labyrinthodonts  from  a  lower  horizon  than 
elsewhere  recorded.  The  discovery  of  Eobaphetes  kansensis  Moodie  in  the  Carbo- 
niferous of  Washington  County  (473)  would  seem  to  indicate  another  labyrinthodont. 
The  tooth  from  Louisville  was  possibly  not  the  first  evidence  of  labyrintho- 
donts in  North  America,  since  the  discovery  by  Marsh  of  Eosaurus  acadianus  from 
the  Coal  Measures  of  Nova  Scotia,  possibly  a  member  of  the  Stereospondylia,  ante- 
dates this  discovery  30  years.  The  specimen  is  preserved  in  the  Museum  at  the 
University  of  Kansas. 

AMPHIBIAN  FOOTPRINTS  FROM  THE  COAL  MEASURES. 

Footprints  may  be  said  to  be  fairly  common  in  the  Coal  Measures  of  North 
America.  Especial  attention  has  been  given  to  the  classification  of  these  objects 
by  G.  F.  Matthew  (408-413),  Dawson  (208-210)  and  others.  Hay  (317,  pp.  538- 
553)  has  given  a  catalogue  of  all  the  species  described  from  the  Coal  Measures  of 
North  America,  to  which  the  reader  is  referred  for  further  information  in  regard  to 
these  interesting  evidences  of  former  animal  activities.  The  writer  has  not  been 
interested  in  the  taxonomy  of  footprints,  but  has  studied  such  as  have  come  to  his 
notice  (465).  A  description  of  the  species  Dromopus  agilis  Marsh  (fig.  43)  is 
given  here,  because  there  is  a  large  slab  in  the  University  of  Kansas  Museum  which 
has  not  been  figured.  Since  the  chief  interest  in  the  present  contribution  is  mor- 
phology, footprints  are  thus  scantily  dealt  with.  Leidy  (374) ,  Dawson  (207) ,  Moodie 
(465,  pi.  lxiv,  fig.  1)  and  others  have  given  various  brief  descriptions  of  Coal  Meas- 
ures footprints,  probably  all  of  which  are  evidences  of  Amphibia  which  are  other- 
wise unknown. 

Dromopus  agilis  Marsh. 

Marsh,  Jour.  Sci.  (3),  xlviii,  p.  82,  pi.  ii,  fig.  3;  pi.  iii,  fig.  3,  1894- 
Hay,  Bull.  U.  S.  Geol.  Surv.,  No.  179,  p.  543,  1902. 

Type:  Specimen  in  the  Yale  University  Museum. 

Horizon  and  locality:  Osage  limestone  (Coal  Measures),  near  Osage  City, 
Kansas. 

In  1894  Professor  Marsh  described  a  collection  of  footprints  which  he  had  secured 
from  Professor  B.  F.  Mudge,  of  Manhattan,  Kansas,  who  had  collected  them  in 
Osage  County,  Kansas,  in  a  rock  quarry,  having  purchased  a  large  quantity  of 
rock  from  the  quarrymen  for  that  purpose.    A  preliminary  note  by  Mudge  (490) 


200 


THE   COAL    MEASURES    AMPHIBIA   OF    NORTH   AMERICA. 


was  published  in  the  Transactions  of  the  Kansas  Academy  of  Science  and  later  copied 
in  the  American  Journal  of  Science.  Professor  Marsh's  description  (406)  of  the 
remains  is  as  follows: 

"  The  impressions  are  well  preserved  in  a  calcareous  shale,  which  separates  readily 
into  thin  slabs,  each  representing  a  surface  of  the  beach  at  the  time  the  footprints  were 
made  upon  it.  A  few  shells  in  the  shale  are  sufficient  to  prove  that  the  formation  is  marine 
(no  shells  are  evident  in  the  slab  at  the  Museum  of  the  University  of  Kansas,  but  the  slab 
is  quite  arenaceous).  Trails  of  annelids,  or  perhaps  of  other  invertebrates,  are  seen  on 
some  of  the  surfaces.  The  footprints  of  vertebrate  animals,  however,  are  of  paramount 
importance,  and  the  large  number  and  variety  of  these  here  recorded  on  a  single  surface, 
if  they  could  be  rightly  interpreted,  would  form  an  interesting  chapter  of  land  vertebrate 
life  in  the  Carboniferous,  about  which  so  little  is  at  present  known." 

Professor  Marsh's  description  of  Dromopus  agilis  is  as  follows: 

"  The  third  series  of  footprints  is  of  special  interest,  and  indicates  an  animal  very 
distinct  from  the  two  already 
described.  The  diagram  rep- 
resents the  impression  of  the 
phalanges  sufficiently  in  de- 
tail to  indicate  (406)  their 
number  and  general  form.  A 
striking  feature  in  the  fore 
and  hind  feet  of  this  animal 
was  the  long,  slender  digits 
terminated  by  sharp  claws. 
Another  point  of  interest,  as 
recorded  in  the  footprints,  is 
that  the  animal  in  walking 
swung  the  hind  feet  outward, 
and  so  near  the  ground  that 
the  ends  of  the  longer  toes 
sometimes    made    trails    in 

the  mud,  marking  accurately  FlG  43._Footprints  of  Promopus  agilis  Marsh,  from  the  Coal  Measures  of 
the  Sweep  Of  the  foot.      This  Osage  County,  Kansas.    Original  slab  in  University  of  Kansas  Museum. 

would    seem   to    indicate    a         Greatly  reduced. 

comparatively  short  hind  leg,  rather  than  the  long,  slender  one  which  the  footprints 

themselves  naturally  suggest. 

"  The  animal  which  made  these  interesting  footprints  was  probably  a  Lacertilian 
rather  than  an  Amphibian,  but  there  is  also  a  possibility  that  it  was  a  primitive  Dinosaur." 

Further  on  Professor  Marsh  remarks  (p.  84) : 

"  So  far  as  at  present  known,  land  vertebrate  life  began  in  the  Carboniferous  age, 
no  footprints  of  other  remains  of  this  kind  having  been  detected  below  the  Subcarbo- 
niferous.  That  such  remains  will  eventually  be  found  in  the  Devonian,  there  can  be  no 
reasonable  doubt,  and  perhaps  even  in  the  Silurian,  if  the  land  surfaces  then  existing 
can  be  explored." 

This  last  statement  of  Marsh's  was,  of  course,  partly  demonstrated  by  the  dis- 
covery of  footprints  in  the  Devonian  rocks  of  Pennsylvania,  which  he  described  in 
1896  as  Thinopus  aiitiquus.     The  footprints  of  Dromopus  agilis  Marsh  which  are 


THE    STEREOSPONDYLOUS    AMPHIBIA. 


201 


preserved  in  obverse  in  the  University  of  Kansas  Museum  are  of  considerably 
greater  length  than  those  described  by  Professor  Marsh.  The  measurements  of  one 
of  the  larger  impressions  are  appended.  There  appear  to  be  series  of  footprints  of 
two  different  animals  preserved  on  the  large  slab  (5  by  7  feet),  but  their  nature  is 
essentially  the  same. 

Measurements  of  Large  Footprint. 

mm.  mm. 

Greatest  length 200      Length  of  shortest  toe 65 

Greatest  width 90      Width  across  heel 50 

Length  of  longest  toe 120 

The  slab  is  No.  5  of  the  University  of  Kansas  Museum  of  Natural  History, 
collected  in  1873  by  Professor  B.  F.  Mudge  at  Osage,  Kansas,  and  presented  by 
him  to  the  museum  in  1875. 

The  following  list  of  Carboniferous  amphibian  (?)  footprints  is  compiled  from 
Hay's  Catalogue  of  Fossil  Vertebrates  of  North  America.  It  is  given  here  for  the 
sake  of  completeness.  Three  types  of  amphibian  footprints  are  described  in  the 
body  of  this  work  and  they  constitute  the  kind  of  material  which  an  ichnologist  has 
for  the  basis  of  his  conclusions.  The  material  is  not  very  satisfactory  for  the 
morphologist,  though  much  can  be  determined  as  to  the  foot  structure. 


Genus  and  species. 


Allopus  littoralis 

Anomccpus  ?  culbertsonii 

gallinidoides 

Anlhracopus  ellangowensis 

Baropus  tentus 

Batrachichnus  plainvillensis. . . . 
Clieirotherium  ?  heterodactylum. 

reiteri 

Collettosaurus  indiantensis 

Crucipes  parvus 

Dromopus  agilis 

Hylopus  caudifer 

hardtngi 

logani 


minor 

trifidus 

sp.  indet 

Limnopus  vagus 

Xanopus  caudutus 

Notalacerta  missouriensis .  . 

Nolamphibia  magna 

Palaosauropus  anliquior. . 

primcevus .  . 

sydenensis . 

unguifer.  .  . 


Thenaropiis  leptodactylus .  .  . 
ovoidaetylus .  .  .  . 
pachydaityltts .  . 
spharodactylus . 


Author. 


Horizon. 


Locality. 


Marsh Coal  Measures. . . 

King Do 

King Do 

Leidy Do 

Marsh Do 

Woodworth   Do 

Do 

Moore Do 

Cox Do 

Butts Do 

Marsh Do 

Dawson Do 

Dawson Do 

Dawson Do 

Dawson Do 

Dawson Do 

Do 

Marsh Do 

Marsh Do 

Butts Do 

Butts Do. 

Dawson. . .  .    Subcarboniferous 
Lea Coal  Measures.. . 

Do 

Carboniferous 
(millstone  grit). 

Coal  Measures.. . 

Do 

Do 

Do 


Dawson. 
Dawson . 


King. 
King. 
King. 
King. 


Osage  County,  Kansas. 
Pennsylvania. 

Do. 

Do. 
Osage  County,  Kansas. 
Massachusetts. 
Pennsylvania. 

Do. 
Indiana. 
Missouri. 

Osage  County,  Kansas. 
Nova  Scotia. 

Do. 

Do. 

Do. 

Do. 

Do. 
Osage  County,  Kansas. 

Do. 
Missouri. 

Do. 
Nova  Scotia. 
Pennsylvania. 
Cape  Breton  Island. 
Nova  Scotia. 

Pennsylvania. 

Do. 
Do. 

Do. 


BIBLIOGRAPHY  OF  THE  FOSSIL  AMPHIBIA,   WITH    ESPECIAL  REFERENCE  TO  THE 
AMPHIBIA  FROM  THE  COAL  MEASURES  OF  NORTH  AMERICA. 

The  following  literature  list,  containing  more  than  six  hundred  titles  ( 1 824-1916), 
is  believed  to  be  fairly  complete.  It  certainly  includes  all  of  the  important 
contributions  to  the  subject  of  the  fossil  Amphibia.  A  few  brief  bibliographies  of 
fossil  Amphibia  have  appeared  from  time  to  time,  in  special  memoirs  on  various 
faunas,  such  as:  Broili  (58),  Schwarz  (541),  Fraas  (242),  von  Ammon  (7),  and  Case 
(98).  None  of  these  have  attempted  a  complete  survey  of  the  field.  The  references 
given  below  have  all,  or  nearly  all,  been  compiled  from  the  original  sources  and  every 
effort  has  been  made  to  have  them  complete  and  accurate. 


1.  Abel,    O.  1912.  Grundziige   der   Paleobiologie   der 

Wirbelthiere.     211-220,  figs.  144-150. 

2.  Agassiz,   L.  1862.  Highly  interesting  discovery  of 

new  Sauroid  remains.  Amer.  Jour.  Sci.  and  Arts, 
Jan.,  xxxiii,  138. 

3.  Alberti,  Fr.  v.  1834.  Beitrag  zu  einer  Monographie 

des  bunten  Sandsteins,  Muschelkalks  und  Keuper. 

4.  .  1864.  Ueberblick  ueber  die  Trias.    235-241. 

5.  Albrecht,  P.  1883.  Note  sur  le  Basioccipital  des 

Batraciens  Anoures.  Bull,  de  Musee  Roy.  d'Hist. 
Nat.  de  Belgique,  n,  195-198,  with  figs,  and  a 
plate. 

6.  American  Naturalist.  1878.  A  new  fauna,     xn, 

327-328  (Ed.). 

7.  Ammon,  Ludwig    von.    1889.  Die  permischen  Am- 

phibien  der  Rheinpfalz,  Munich.  1-117,  pis.  1-5, 
with  extensive  bibliography. 

8.  Andrews,  C.  W.  1895.  Note  on  a  specimen  of  Kera- 

terpeton  galvani  Huxley,  from  Staffordshire.  Geol. 
Mag.,  Dec,  iv,  n,  81-84,  fig. 

9.  Arldt,  Theodor.  1907.  Die  Entwicklung  der  Konti- 

nente  und  ihre  Lebewelt.    333,  Karte  10. 

10.  .  1909.  Die  Stegocephalen  und  ihre  Stellung 

unter  den  Wirbelthieren.  Naturw.  Rundschau, 
xxiv,  353-355- 

11.  Atthey,  T.  1876.  On  Anthracosaurus  russelli.    Ann. 

and  Mag.  Nat.  Hist.,  Ser.  4,  xvm,  146-167,  pis. 
8-1  i. 

12.  Bailey,  W.  H.  1866.  On  the  new  discovery  of  fossil 

reptiles  in  the  Carboniferous  of  southern  Ireland. 
Geol.  Mag.,  11,  No.  20,  84. 

1875.  Description  of  a  new  species  of  Laby- 


13 


14- 


15- 
10. 


17. 


rinthodont  Amphibian  from  the  Coal  at  Jarrow 

Colliery  near  Castlecomer,  Kilkenny.    Rept.  Brit. 

Assn.  Adv.  Sci.,  62.     (Describes  Anthracosaurus 

edgei.) 
— ■ — ■.   1884.  Some  additional  notes  on  Anthraco- 
saurus edgei  Bailey.    Rept.  Brit.  Assn.  Adv.  Sci., 

96-97  (1883). 
Barkas,  T.  P.   1868.  On  the  fauna  of  the  low  main 

Coal  Seam,  Northumberland.    Geol.  Mag.,  v,  580. 
.  1869.  Unusual     forms    of    Ctenoptychius. 

Geol.  Mag.,  Jan.,  vi,  43,  2  woodcuts. 
.   1869.  On  a  supposed  mammalian  tooth  from 

the  Coal  Measures.     Monthly  Micros.  Jour.,  11, 

104,  figs.  1-5. 


18.  Barkas,  T.  P.  1869.  On  the  discovery  of  a  molar  of 
a  large  reptile  in  the  Northumberland  Coal  Meas- 
ures. Ann.  and  Mag.  Nat.  Hist.,  m,  419.  (Pter- 
oplax  cornuta.) 

19. .     1869.    Reptile    Remains  and  Climadoxus. 

Ann.  and  Mag.  Nat.  Hist.,  iv,  438-440. 

20.  .  1873.  A  manual  of  Coal  Measure  paleon- 
tology.   London. 

21.  Barrell,  Joseph.  1906.  Origin  and  significance  of 

Mauch   Chunk  shale.     Bull.   Geol.   Soc.   Amer., 
xvm,  460. 

22.  Baur,  George.   1886.  Die  alteste  Tarsus  (Archego- 

saurus).    Zool.  Anz.,  No.  216,  9  Jahrg.,  104-106. 

23. .  1886.  The  oldest  Tarsus   (Archegosaurus). 

Amer.  Nat.,  xx,  173-174. 

1886.  Ueber  die  Homologien  einiger  Schadel- 


24 


25- 


26. 


27- 


28. 


29. 


30- 


31- 


32. 


32a 


33- 


knochen  der  Stegocephalen  und  Reptilien.    Anat. 

Anz.,  1,  No.  13,  348-350. 
.   1886.  Ueber  die   Morphologie   der  Wirbel- 

saule  der  Amnioten.    Biol.  Centralb.,  vi,  Nr.  1 1-12, 

322-342,  353-363- 
.  1887.  On  the  morphology  of  ribs.     Amer. 

Nat.,  Oct.,  942-945  (Archegosaurus). 
.  1887.  Nachtriigliche   Notiz  zu  meinen   Be- 

merkungen    ueber    die    Homologie    der    Schadel- 

knochen  der  Stegocephalen  und  Reptilien.    Anat. 

Anz.,  2,  No.  21,  657-658. 
.  1888.  Morphogenie  der  Carpus  und  Tarsus 

der  Wirbelthiere.    Pt.  I,  Batrachia.    Jena,  Verlag 

von  Gustav  Fischer,  i-88,  pis.  1-3,  figs. 
.   1894.  Bemerkungen  liber  die  Osteologie  der 

Schliifengegend  der  hoherer  Wirbelthiere.    Anat. 

Anz.,  x,  315-330. 
.  1896.  Bemerkungen  iiber  die  Phylogenie  der 

Schildkroten.     Anat.  Anz.,  xn,   561-570.     (Dis- 

sorophus.) 
.  1896.  The    Stegocephali.     A    phylogenetic 

study.    Anat.  Anz.,  20ten  Marz,  xi,  No.  22,  657- 

673,  with  bibliography  and  eight  figures. 
— .  1897.  Archegosaurus.     Amer.   Nat.,   xxxi, 


975-980. 
.    1897.   Ueber  die  systematische  Stellung  der 

Microsaurier.     Anat.  Anz.,  xiv,  148-151. 
Bayer,     Franz.   1880.  Palaeobatrachus    bohemicus 

von  Meyer,  aus  der  Braunkohle  von  Freudenhain. 

Sitzber.  d.  k.  bohm.  Ges.  Prag,,  291-298,  1  taf. 


BIBLIOGRAPHY   OF   THE   FOSSIL   AMPHIBIA. 


203 


34.  Beasley,  H.  C.  1900.  Notes  on  the  type  specimen 

of  Cheirotherium  herculis  Egerton.  Proc.  Liver- 
pool Geol.  Soc.,  xlii,  81,  pi.  v. 

35.  Bernard,  Felix.  1895.  Elements  de  Paleontologie. 

Batraciens,  740-760,  with  figs. 

36.  Beyrich,  Ernst.     1850.    Ueber  einige  organische 

Reste  der  Lettenkohlenbildung  in  Thueringen. 
Zeit.  d.  deutsch.  geol.  Gesell.,  11,  165. 

37.  Bieber,  V.  1880.  Ueber  zwei  neue  Batrachier  der 

Bohm.  Braunkohlenformation.  Sitz.  d.  k.  Akad. 
d.  Wissen.,  Wien.,  lxxxii,  abth.  1,  117,  taf.  iii, 
fig.  1. 

38.  Binney,  E.  W.   1883.  Trans.  Manchester  Geol.  Soc, 

vi,  p.  42. 

39.  Blanchard,  R.  1 88 1.  Sur  les  Glandes  cloacale  et 

Pelvien  et  sur  la  Papille  cloacale  des  Batraciens 
urodeles.    Zool.  Anz.,  iv,  9,  34. 

40.  Blanford,  W.  T.  1884.  Address  to  the  geological 

section  of  the  British  Association  at  Montreal, 
21  pp.;  also  Nature,  xxx. 

41.  Bolkay,  S.  J.  de.  191 1.  On  a  Pleistocene  predecessor 

of  Rana  fusca  Ros.  Mittheil.  aus  d.  Jahrb.  k.  Ung. 
Geol.  R.,  xix,  H.  3,  155-160,  figs.  1-7. 

42.  Bolton,  H.  1896.  The  animal  life  of  the  Lancashire 

Coal  Measures.  Trans.  Manchester  Micros.  Soc, 
123-135  (1895)- 

43-  .  1904.  The  paleontology  of  the  Lancashire 

Coal  Measures.  Trans.  Manchester  Geol.  Soc, 
xxviii,  378-420,  578-650,  668-689. 

44.  Boulenger,  G.  A.  1890.  On  the  presence  of  ptery- 
goid teeth  in  a  tailless  batrachian  (Pelobates  cul- 
tripes),  with  remarks  on  the  localization  of  teeth 
on  the  palate  in  batrachians  and  reptiles.  Proc 
Zool.  Soc.  London,  664-666. 

-.  1904.  On  the  characters  and  affinities  of  the 

Triassic  reptile  Telerpeton  eligense.  Proc  Zool. 
Soc.  London,  470-480,  pi.  xxx. 

1910.  Les  Batraciens  et  principalement  ceux 


45- 


46. 


d'Europe.    1.    Paris. 

47.  Bradley,  Frank    H.    1870.    Geology  of  Grundy 

County.    Geol.  Surv.  Illinois,  iv,  196. 

48.  Branco,  W.  1887.  Weissia  bavarica,  nov.  gen.  et 

sp.  von  Stegocephalia,  ein  neuer  Stegocephale  aus 
dem  unteren  Rothliegenden.  Jahrb.  d.  k.  Preuss. 
Geol.  Landesanstalt  f.  1886,  22-39,  ta^-  ';  a's° 
Neues  Jahrb. f.Mineral., Geol. u.Paleon.,  1888, 117. 

49.  Branson,  E.  B.  1905.  Structure  and  relationships  of 

American  Labyrinthodontidje.  Jour.  Geol.,  xin, 
No.  7,  568-610,  figs.  1-19. 

50.  .  1910.  Amphibian  footprints  from  the  Mis- 

sissippian  of  Virginia.  Jour.  Geol.,  xvin,  No.  6, 
356-358,  1  fig- 

51.  Braun,  M.  von.  1 841.  Bericht  tiber  die  Versamm- 

lung  deutscher  Naturforscher  und  Aertze.  Braun- 
schweig, 74-75.     (Trematosaurus.) 

52.  Braun,  M.  1878.  Ueber  aussere  Hilfsorgane  bei  der 

Begattung  von  Triton  viridescens.  Zool.  Anz.,  I, 
124. 

53.  British  Museum.  1905.  A  guide  to  the  fossil  rep- 

tiles, amphibians  and  fishes  in  the  department  of 
geology  and  paleontology  in  the  British  Museum 
(Natural  History).     8  plates,  116  figs.,  8th  ed. 

54.  Brodie,  P.  B.  1859.  On  the  occurrence  of  footsteps 

of  Cheirotherium  in  the  Upper  Keuper  of  War- 
wickshire.   Quart.  Jour.  Geo!.  Soc,  xvi,  278. 

55.  Broili,   F.  1899.  Ein    Beitrag  zur   Kenntniss  von 

Eryops  megacephalus  Cope.  Paleontographica, 
xlvi,  61-84,  pis.  8-10. 


56. 

57- 
58. 
59- 
60. 

61. 

62. 

63- 

64. 
65- 

66.  - 


Broili,  F.  1902.  Beitragc  zur  Kenntniss  von  Diplo- 
caulus  Cope  (D.  magnicornis).  (Paleonto- 
graphica, li,  8.)  Vorlaufigc  Mittheilung,  Cen- 
tralb.  f.  Mineral.,  Geol.  u.  Paleon.,  536-541,  with 
figs. 

.  1904.  Stammreptilicn.     Anat.    Anz.,    xxv, 

No-  23,  577-587- 

— .  1904.  Permische  Stegocephalen   und   Rep- 

tilien.    Paleontographica,  li,  1-120,  5  pis. 

.  1905.  Beobachtungen  an  Cochleosaurus  bc- 

hemicus.    Paleontographica,  L,  1-16,  pis.  1-2. 

.  1906.    Ein    Stegocephalen    Rest    aus    den 

Bayrischcn  Alpen.    Centralb.  f.  Mineral.,  Geol.  u. 
Paleon.,  No.  18,  568-571. 

1908.  Ueber   die   Rachitomen   Wirbel   der 


Stegocephalen.    Monatsberichten  d.  deutsch.  geol. 

Gesell.,  60,  Nr.  8/10,  235-240,  figs.  1-7. 
•  1908.  Ueber   Sclerocephalus  aus  der  Gas- 

kohle  von  Nurschan  und  das  Alter  dieser  Ablager- 

ungen.    Jahrb.  d.  k.  k.  Geol.  Reichsanstalt,  lviii, 

Heft  1,  49-69,  Taf.  1. 
.  1908.  Systematische    und    Biologische   Be- 

merkungen   zu    der    permischen   Gattung   Lyso- 

rophus.    Anat.  Anz.,  xxxm,  Nr.  11/12,  290-298. 
1909.  Reviews  of  Literature.     Neues  Jahrb. 


f.  Mineral.  Geol.  u.  Paleon.,  Bd.  11,  Heft  1,  132-138. 

.  1913.  Ueber   zwei    Stegocephalenreste   aus 

dem  texanischen  Perm.    Neues  Jahrb.  f.  Mineral., 
Geol.  u.  Paleon.,  Bd.  1,  Heft  2,  96-100,  Taf.  IX. 
— .  1913.  Unser  Wissen  iiber  die  altesten  Tetra- 


67. 
68. 

Go- 
To. 
71- 

72. 

73- 

74- 
75- 
76. 

77- 


poden.    Fortschr.  d.  naturwissenschaftlichen  For- 

schung,  Halle,  Bd.  vm,  51-93,  figs.  14-62. 
Bronn,   H.   G.  1852-54.  Lethea  Geognostica,  mit 

Atlas.    Bd.  1  (2),  Kohlen  Periode,  von  F.  Roenier. 
Broom,  R.  1903.   On  a  new  Stegocephalian    (Ba- 

trachosuchus  browni)   from  the  Karoo   Beds  of 

Aliwal   North,  South  Africa.    Geol.    Mag.,  n.  s., 

Dec,  iv,  x,  No.  473-499.  1-2.  figs- 
.  1904.  On  a  new  South  African  Labyrinthc- 

dont     (Cyclosaurus   albertyni).     Records  of  the 

Albany  Museum,  1,  No.  3. 

-,  1907.  On    the   geological   horizons   of   the 


vertebrate    genera    of     the    Karoo    Formation. 
Records  of  the  Albany  Museum,  11,  No.  1. 

-.  1908.  On  a  new  Labyrinthodont   (Rhine- 


suchus  whaitsi)  from  the  Permian  beds  of  South 
Africa.  Ann.  So.  African  Museum,  iv,  pt.  viii, 
May  1908,  373. 

-.  On  the  nomenclature  of  the  elements  of  the 


amphibian  shoulder  girdle. 

-.  1910.  A  comparison  of  the  Permian  reptiles 


of  North  America  with  those  of  South  Africa. 
Bull.  Amer.  Mus.  Nat.  Hist.,  xxvin,  art.  xx,  197- 

234- 

191 3.  On  the  structure  of  the  mandible  in 


the  Stegocephalia.    Anat.  Anz.,  Bd.  45,  No.  2/3, 
73-78,  figs. 

19 1 3.  Studies  on  the  Permian  temnospondy- 


lous  Stegocephalians  of  North  America.    Bull.  Am. 

Mus.  Nat.  Hist.,  xxxn,  563-595,  21  figs. 
Brown,    Barnum.    1908.      The  Conard  fissure,  a 

Pleistocene  bone  deposit  in   northern  Arkansas, 

with  description  of  two  new  genera  and  twenty 

new  species  of  mammals.    Mem.  Amer.  Mus.  Nat. 

Hist.,  ix,  pt.  iv,  206,  pi.  xxii. 
Browne,  M.  1893.  On  some  vertebrate  remains  not 

hitherto  recorded  from  the  Rhaetic  beds  of  Britain. 

Rept.  Brit.  Assn.  Adv.  Sci.,  748. 


204 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


78.  Brownrigg,  W.  B.   1865.  Jour.  Roy.  Geog.  Soc. 

Ireland,  I,  pt.  2,  145. 

79.  Budgett,  J.  S.  1902.  On  the  structure  of  the  larval 

Polypterus.  Trans.  Zool.  Soc.  London,  xvi,  pt. 
vil,  Oct.;  also  Works,  154-176,  pis.  x-xii,  1907. 

80.  Burmeister,   H.  1840.  Die  Labyrinthodonten  aus 

dem  bunten  Sandstein  von  Bernburg.  4  plates 
(same  1849)  (?). 

81. .  1850.  Die  Labyrinthodonten  aus  dem  Saar- 

briicken  Steinkohlengebirge.    Quarto,  pis.  1-4. 

82.  Butts,    Edward.  1891.  Recently   discovered   foot- 

prints of  the  amphibian  age  in  the  Upper  Coal 
Measure  Group  of  Kansas  City,  Missouri.  Kans. 
City  Scientist,  v,  No.  2,  17-19,  figs. 

83.  .  1891.  Footprints  of  new  species  of  amphib- 
ians in  the  Upper  Coal  Measure  Group  of  Kansas 
City,  Missouri.     Kans.  City  Scientist,  V,  44,  figs. 

84.  Carus,  J.  Victor.  1875.  Labyrinthodontia.    Hand- 

buch  der  Zoologie,  Bd.  I,  485-486. 

85.  Case,  E.  C.   1898.  The  significance  of  certain  changes 

in  the  temporal  region  of  the  primitive  reptilia. 
Amer.  Nat.,  xxxn,  No.  374,  69. 

.  1900.  The  vertebrates  from   the   Permian 

bone  bed  of  Vermilion  County,  Illinois.  Jour. 
Geol.,  viii,  698-729,  5  plates. 

1902.  On  some  vertebrate  fossils  from  the 


86. 


87. 


88. 


89. 


90. 


91- 


92. 


93- 


94. 


95- 


96. 


97- 


98. 


99- 


Permian  beds  of  Oklahoma.  2d  Biennial  Rpt. 
Dept.  Geol.  and  Nat.  Hist.  Territory  Oklahoma, 
1901-1902,  62-68. 

1902.  Paleontological  notes.    Jour.  Geol.,  x, 


256-261. 

-.  1903.  New  or  little-known  vertebrates  from 


the  Permian  of  Texas.    Jour.  Geol.,  xi,  394-402. 
1907.  Additional  description  of  the  genus 


Zatrachys  Cope.     Bull.  Amer.   Mus.    Nat.  Hist., 
xxiii,  art.  xxix,  665-668. 

.  1907.  Revision  of  the  Pelycosauria  of  North 

America.     Pub.  No.  55,  Carnegie  Inst.  Washing- 
ton, Stegocephalia,  159. 

-.  1908.  On  the  value  of  the  evidence  furnished 


by  vertebrate  fossils  of  the  age  of  certain  so-called 
Permian  beds  in  America.  Jour.  Geol.,  xvi,  No.  6, 
572-580. 

.  1908.  Notes  on  the  skull  of  Lysorophus  tri- 

carinatus  Cope.  Bull.  Amer.  Mus.  Nat.  Hist., 
xxiv,  531-533,  fig.  1. 

-.  1908.  Description  of  vertebrate  fossils  from 


the  vicinity  of  Pittsburgh,  Pa.  Annals  of  the  Car- 
negie Museum,  iv,  Nos.  iii-iv,  234-241,  pi.  lix,  Apr. 
1908.  A  great  Permian  delta  and  its  verte- 


brate life,  with  restorations  by  the  author.    Pop. 
Sci.  Month.,  lxxiii,  No.  6,  557-568. 

.  1910.  New   or    little-known    reptiles    and 

amphibians  from  the  Permian?  of  Texas.  Bull. 
Amer.  Mus.  Nat.  Hist.,  xxvm,  art.  xvn,  163-181. 
191 1.  A    revision    of    the    Cotylosauria    of 


North   America.     Pub.  No.   145,  Carnegie    Inst. 

Washington. 
.  191 1.  Revision  of  the  Amphibia  and  Pisces 

of  the  Permian  of  North  America.    Pub.  No.  146, 

Carnegie  Inst.  Washington,  1-148,  pis.   1-25,  51 

text-figs. 
.  1912.  Paleozoic    Reptilia    and    Amphibia. 

Bull.  Geol.  Soc.  Amer.,  23,  200-204. 

1915.  The    Permo-carboniferous    beds    of 


North  America  and  their  vertebrate  fauna.  Pub. 
No.  207,  Carnegie  Inst.  Washington,  1-176,  pis. 
1-24. 


101.  Chamberlin,   T.   C.  1900.  On  the  habitat  of  the 

early  vertebrates.    Jour.  Geol.,  vm,  400-412. 

102.  ,  and    R.    D.    Salisbury.  1906.  Geology, 

earth  history.    Amphibia,  11,  606-610. 

103.  Clark,  C.  T.  1847.  On  the  neighborhood  of  Bom- 

bay and  certain  beds  containing  fossil  frogs. 
Quart.  Jour.  Geol.  Soc.,  in,  221-224.  (See 
Owen,  1847.) 

104.  Colliery   Guardian.  Description   of  claspers   of 

Amphibia.    Mar.  10,  1 87 1.     (Barkas.) 

105.  Cope,  E.  D.  1865.  On  Amphibamus  grandiceps,  a 

new  Batrachian  from  the  Coal  Measures.  Proc. 
Acad.  Nat.  Sci.  Phila.,  134-137.  See  also  Geol. 
Surv.  111.,  11,  135,  fig.,  pi.  32,  1866. 

106.  .  1866.  Remarks  on  extinct  vertebrates  of 

the  Mesozoic  Red  Sandstone.  Proc.  Acad.  Nat. 
Sci.  Phila.,  249.    (Mastodonsaurus  durus.) 

107.  .  1866.   Supplement   to   the   description   of 

vertebrates.  Geol.  Surv.  111.,  n,  135-141,  fig., 
pi.  32,  fig.  8.     (Amphibamus.) 

108.  .  1866.  On  the  structure  and  distribution  of 

the  genera  of  the  arciferous  Anura.  Jour.  Acad. 
Nat.  Sci.  Phila.,  2d  ser.,  v,  67-112. 

109. .  1866.  On  the  families  of  Raniform  Anura. 

Jour.  Acad.  Nat.  Sci.  Phila.,  189. 

no. .  1868.  Synopsis  of  the  extinct  Batrachia  of 

North  America.     Proc.  Acad.  Nat.  Sci.  Phila., 
208-221. 
in.  .  1868.  Two  Batrachians  from  North  Caro- 
lina.   Proc.  Acad.  Nat.  Sci.  Phila.,  xx,  211-212. 
(Pariostegus  myops.) 

.  1869.  Synopsis  of  the  extinct    Batrachia, 

Reptilia,  and  Aves  of  North  America.  Trans. 
Amer.  Phil.  Soc,  xiv,  1-252,  pis.  1-14. 

.  1 87 1.  Observations  on  extinct  Batrachia. 

Proc.  Acad.  Nat.  Sci.  Phila.,  53. 

1 87 1.  Supplement  to  the  synopsis  of  the 


112. 

113- 
114. 

"5- 

116. 
117. 

118. 

119. 


extinct  Batrachia  and  Reptilia  of  North  America. 
Proc.  Amer.  Phil.  Soc.,  xn,  41-52. 

1871.  Observations  on  the  extinct  Batra- 


chian fauna  of  the  Carboniferous  of  Ohio  (Linton). 
Proc.  Amer.  Phil.  Soc,  xn,  177. 
— .    1872.    Carboniferous     reptiles     of     Ohio. 


Amer.  Nat.,  vi,  46. 

.  1873.  Observations  on  the  distribution  of 

certain  extinct  Vertebrata  in  North  Carolina. 
Proc.  Amer.  Phil.  Soc,  xn,  210-216. 

.  1873.  On  some  new  Batrachia  and  Fishes 

from  the  Coal  Measures  of  Linton,  Ohio.  Proc. 
Acad.  Nat.  Sci.  Phila.,  340-343. 

.  1874.  Supplement  to  the  extinct  Batrachia, 

Reptilia  and  Aves  of  North  America.  I.  Cata- 
logue of  the  air-breathing  Vertebrata  from  the  Coal 
Measures  of  Linton,  Ohio.  Trans.  Amer.  Phil. 
Soc,  xv,  261-278. 

1875.  Check-list  of   North  American  Ba- 


trachia and  Reptilia,  with  a  systematic  list  of  the 
higher  groups  and  an  essay  on  geographical 
distribution  based  on  the  specimens  contained  in 
the  U.  S.  National  Museum.  Bull.  U.  S.  Nat. 
Mus.,  1,  7-12. 

.  1875.   Synopsis  of  the   Vertebrata   whose 

remains  have  been  preserved  in  the  formations 
of  North  Carolina.  Appendix  "b"  by  Kerr, 
W.  C.  Rept.  Geol.  Surv.  N.  C,  Raleigh,  29-52, 
pis.  v-viii. 

.  1875.  The  extinct  Batrachia  of  Ohio.    Proc. 

Acad.  Nat.  Sci.  Phila.,  16. 


BIBLIOGRAPHY  OF   THE   FOSSIL  AMPHIBIA. 


205 


123- 
124. 

125- 

126. 
127. 
128. 

129. 

130. 
I3i- 

132. 

133- 
134- 

iss- 
137- 

138. 

139- 
140. 
141. 

142. 

143- 
144. 


Cope,  E.  D.  1875.  Synopsis  of  the  extinct  Batrachia 
from  the  Coal  Measures.  Geol.  Surv.  Ohio,  11, 
pt.  11,  351-410,  pis.  xxvi-xlv. 

.  1876.  On  some  extinct  reptiles  and  Batra- 
chia from  the  Judith  River  and  Fox  Hills  Beds 
of  Montana.  Proc.  Acad.  Nat.  Sci.  Phila.,  357; 
Pal.  Bull.  23. 

•  1876.  Description  of  some  vertebrate  re- 
mains from  the  Fort  Union  Beds  of  Montana. 
Proc.  Acad.  Nat.  Sci.  Phila.,  260.  (Ccratodus 
hicroglyphus,  an  amphibian  clasper?.) 

.  1877.  On   the   fossil   remains   of   Reptilia 

and  fishes  from  Illinois.     Proc.  Acad.  Nat.  Sci. 
Phila.,  405. 

1877.  On  the  Vertebrata  of  the  Bone  Bed 


of  eastern  Illinois.    Proc.  Amcr.  Phil.  Soc,  xvn, 

53-64- 
.  1877.  Report  on  the  geology  of  the  region 

of  the  Judith  River,  Montana,  and  on  vertebrate 

fossils  obtained  on  or  near  the  Missouri  River. 

Bull.  U.  S.  Geol.  and  Geog.  Surv.  of  Territories, 

hi,  art.  xix,  Batrachia,  573,  576. 
.  1877.  Descriptions  of  Vertebrata  from  the 

Permian  and  Triassic  Formations.    Proc.  Amer. 

Phil.  Soc.,  xvir,  182-193. 
.  1877.  Extinct     Vertebrata     from     North 

Carolina.    Proc.  Amer.  Phil.  Soc,  17,  182. 
.  1877.  A  continuation  of  researches  among 

the  Batrachia  of  the  Coal  Measures  of  Ohio. 

Proc.  Amcr.  Phil.  Soc,  xvi,  573-578. 
.  1878.  Description  of  extinct  Batrachia  and 

Reptilia  from  the  Permian  of  Texas.    Proc.  Amer. 

Phil.  Soc,  xvii,  522-526. 

1878.  The  vertebra  of  Rachitomus.    Amer. 


Nat.,  xii,  633.     (Eryops.) 

1879.  The    relations   of    the    horizons    of 


extinct  Vertebrata  of  Europe  and  North  America. 
Bull.  U.  S.  Geol.  and  Geog.  Surv.  of  the  Terri- 
tories, v,  No.  1,  33-34. 

-.  1880.  Extinct  Batrachia.    Amer.  Nat.,  xiv, 


609-610.     Reviews  Fritsch's  and  Wiedersheim's 

papers. 
.     1880.     The     structure    of    the   Permian 

Ganocephala.       Amer.     Nat.,      xiv,     383-384. 

(Eryops.) 
.  1 88 1.  The    Permian    Formation    of   New 

Mexico.     Amer.  Nat.,  xv,  1020-1021.     (Eryops 

and  Zatrachys.) 
.  1 88 1.  Catalogue    of    Vertebrata    of    the 

Permian  Formation  of  the  United  States.    Amer. 

Nat.,  xv,  162-164. 
— ■.   1 88 1.  Ein  Uebergangsglied  von  den  Am- 

phibicn  zu  den  Reptilien  (Cricotus).    Auszug  in 

Kosmos  von  Krause,  Bd.  9,  230-231. 
.  1 88 1.  On  some  new  Batrachia  and  Reptilia 

from  the  Permian  Beds  of  Texas.     Bull.  U.  S. 

Geol.  Surv.  Terr.,  vi,  art.  II,  79-82. 

-.  1882.  Third  contribution  to  the  history  of 


the  Vertebrata  of  the  Permian  Formation  of 
Texas.  Proc.  Amer.  Phil.  Soc,  xx,  447-461 ;  Pal. 
Bull.  35,  451. 

1882.     The    Rachitomous  Stegocephalia. 


Amer.  Nat.,  xvi,  335. 
.     1882.     Permian  Vertebrata.    Amer.  Nat., 


xvi,  925. 

.  1883.  Fourth   contribution   to  the  history 

of  the  Permian  Formation  of  Texas.    Proc.  Amer. 
Phil.  Soc,  xx,  628-636. 


145.  Cope.E.D.  1883.  On  some  Vertebrata  from  the  Per- 

mian of  Illinois.     Proc.  Acad.  Nat.  Sci.  Phila., 
108-110. 

146.  .  1884.  The     Batrachia    of     the     Permian 

Period  of  North  America.     Amer.   Nat.,  xvm, 
26-39,  pis.  ii-v,  7  figs. 

1 884.  Fifth  contribution  to  the  fauna  of  the 


147- 


148. 


149. 


150. 


151- 


152. 


'53- 


154- 


155- 


156. 


'57- 
158. 

«59- 
160. 
161. 

162. 

163. 
164. 

165. 

166. 
167. 

168. 
169. 


Permian  of  Texas  and  Indian  Territory.     Proc. 
Amer.  Phil.  Soc,  xxn,  28-30,  pi.  1 ;  Pal.  Bull.  39. 
.  1884.  Note  on  the  phylogeny  of  the  Verte- 
brata.   Amer.  Nat.,  xvm,  1255-1256. 

1885.  The  Batrachia  of  the  Permian  Beds 


of  Bohemia  and  the  Labyrinthodon  from  the 
Bijori  Group,  India.  Amer.  Nat.,  xix,  592-594. 
1885.  Paleontological  nomenclature.    Geol. 


Mag.,  London  (3),  11,  575.     (Batrachia  for  Am- 
phibia.) 

.  1885.  The  Batrachia  of  the  Permian  Beds 

of  Bohemia.    Geol.  Mag.,  Ixindon  (3),  n,  527. 

.  1885.  Second   continuation   of   researches 

among  the  Batrachia  of  the  Coal  Measures  of 
Ohio.    Proc.  Amer.  Phil.  Soc,  xxn,  405-408. 

.  1885.  On  the  evolution  of  the  Vertebrata, 

progressive  and  retrogressive.   Amcr.  Nat.,  xix, 
244,  Mar. 

1886.  On   the  structure  and  affinities  of 


the  Amphiumidx.    Proc.  Amer.  Phil.  Soc,  xxm, 
No.  123,  442-445.  July- 

1886.  The  Batrachian  intcrcentrum.  Amer. 


Nat.,  xx,  76,  175. 
.  1886.  Systematic  catalogue  of  the  species 

of  vertebrates  found  in  the  Beds  of  the  Permian 

Epoch  of  North  America.     Trans.  Amer.  Phil. 

Soc,  xvi,  285-296,  pis.  ii-iii. 
.  1887.  A  contribution  to  the  history  of  the 

Vertebrata  of  the  Trias  of  North  America.    Proc. 

Amcr.  Phil.  Soc,  xxiv,  209.    (Eupclor  durus.) 
— .  1888.  On  the  intercentrum  of  terrestrial 


vertebrates.    Trans.  Amcr.  Phil.  Soc,  xvi,  243- 
253,  pl-  1,  6  figs. 

1888.  The  Ossicula  Auditus  of  the  Batra- 


chia.   Amer.  Nat.,  xxn,  464-467,  pl.  vi. 

1888.  The  pineal  eye  in  extinct  vertebrates. 


Amer.  Nat.,  xxn,  914-917,  pis.  xv-xviii. 

1888.  On  the  shoulder  girdle  and  extremi- 


ties of  Eryops.     Trans.  Amer.  Phil.  Soc,  xvi, 
362-367,  with  plate. 

1888.  On  the  relation  of  the  hyoid  and 


otic  elements  of  the  skeleton  in  the  Batrachia. 
Jour.  Morphol.,  11,  297-310. 

.  1889.  The  Batrachia  of  North  America.  Bull. 

U.  S.  Nat.  Mus.,  No.  34,  i-vi,  7-514,  pis.  i-lxxxvi. 

.  1889.  Synopsis  of  the  families  of  Verte- 
brata. Amer.  Nat.,  xxm,  849-877.  (Published 
in  1898  as  "Syllabus  of  Lectures.") 

-.  1892.  A  preliminary  report  on  the  verte- 


brate paleontology  of  the  Llano  Estacado.    Geol. 
Surv.  Texas,  4th  Ann.  Rept.,  12-17. 

1892.  On   the   phylogeny   of   Vertebrata. 


Proc.  Amer.  Phil.  Soc,  xxx,  278-280. 

.  1892.  On  some  new  and  little-known  Pal- 
eozoic vertebrates.  Proc.  Amcr.  Phil.  Soc,  xxx, 
221-229,  pls-  vii-viii. 

.  1893.  Fritsch's  Fauna  of  the  Gaskohle  of 

Bohemia.     Amcr.  Nat.,  xxyii,  709. 

.  1895.  Some  new  Batrachia  from  the  Per- 
mian Beds  of  Texas.  Proc.  Amer.  Phil.  Soc, 
.xxxiv,  452-457,  plate  ix,  Nov.  15. 


206 


THE   COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


170. 
171. 

172. 

173- 
174. 

175- 

176. 

177. 

178. 
179- 

180. 


Cope,  E.  D.  1895.  A  Batra  chianarmadillo.  Amer. 
Nat.,  xxix,  998,  Nov. 

.  1 896.  The  Paleozoic  reptilian  order  Cotylo- 

sauria.  Amer.  Nat.,  xxx,  303.  (Describes 
Eosauravus  copei  Williston,  oldest  known  reptile.) 
1896.  Permian  land  vertebrates  with  cara- 


paces.   Amer.  Nat.,  xxx,  936,  pi.  xxi. 

-.  1896.  The   ancestry   of   the   Testudinata. 


Amer.  Nat.,  xxx,  399. 
.  1896.  Appendix  on  a  species  of  Trimero- 

rhachis.    Proc.  Amer.  Phil.  Soc,  xxxv,  137. 
.   1896.   The   reptilian   order   Cotylosauria. 

Supplement:     Some    new    Batrachia    from    the 

Permian  beds  of  Texas.    Proc.  Amer.  Phil.  Soc, 

xxxiv,  455. 

1897.  On  new   Paleozoic  Vertebrata  from 


183. 


184. 


185. 


t86. 


187. 


Illinois,  Ohio,  and  Pennsylvania.  Proc.  Amer. 
Phil.  Soc,  xxxvi,  71-90,  three  plates. 

.  1898.  Syllabus  of  Lectures  on  the  Verte- 
brata. Batrachia,  43-51.  Published  for  Uni- 
versity of  Pennsylvania. 

Cox,  E.  T.  1874.  Colettosaurus  indianaensis.  5th 
Ann.  Rept.  Geol.  Surv.  Indiana,  247,  pi.  1. 

Credner,  Hermann.  1881.  Ueber  Branchiosaurus 
amblystomus,  einen  neuen  Stegocephalen  aus  den 
Rothliegenden-Kalke  von  Niederhasslich  im 
Plauen'schen  Griinde.  Sitzungsb.  naturf.  Gesell. 
zu  Leipzig,  43. 

.  1 88 1.  Die  Stegocephalen  und  Saurier  aus 

dem  Rothliegenden  des  Plauen'schen  Grundes  bei 
Dresden.  I  Theil:  Einleitung  und  Branchio- 
saurus gracilis.  Zeit.  d.  deutsch.  geol.  Gesell., 
xxxm,  298-329,  taf.  xv-xviii. 

.  1 88 1.  Die  Stegocephalen  und  Saurier,  etc. 

II  Theil:  Branchiosaurus  amblystomus.  Zeit. 
d.  deutsch.  geol.  Gesell.,  xxxm,  574-603,  taf. 
xxii-xxiv. 

.  1881.  Ueber  einige  Stegocephalen  (Laby- 

rinthodonten)  aus  dem  sachsischen  Rothliegen- 
den. Sitzungsb.  naturf.  Gesell.  Leipzig,  1-11. 
(Note  on  Branchiosaurus  and  Melanerpeton,  43, 
45-) 

.  1882.  Ueber  Melanerpeton  Fritsch  aus  dem 

Rothliegenden-Kalk  von  Niederhasslich  im  Plau- 
en'schen Grundes.  Sitzungsb.  naturf.  Gesell. 
Leipzig,  45-47- 

.  1882.  Die  Stegocephalen  und  Saurier,  etc. 

III  Theil:  Pelosaurus  laticeps,  Archegosaurus 
decheni,  A.  latirostris.  Zeit.  d.  deutsch.  geol. 
Gesell.,  xxxi v,  213-237,  tafeln  xii-xiii. 

.  1883.  Die  Stegocephalen  und  Saurier,  etc. 

IV  Theil:  Branchiosaurus  gracilis,  Acanthos- 
toma  vorax,  Melanerpeton,  Discosaurus  permi- 
anus.  Zeit.  d.  deutsch.  geol.  Gesell.,  xxxv,  275- 
300,  tafeln  xi-xii. 

.  1885.  Die  Stegocephalen  aus   dem   Roth- 
liegenden des  Plauen'schen  Grundes  bei  Dresden. 

V  Theil:  Melanerpeton  pulcherrimum,  Pelo- 
saurus laticeps,  Archegosaurus;  Sparagmites 
arciger,  Hylonomus  fritschii,  Hyloplesion.  Zeit. 
d.  deutsch.  geol.  Gesell.,  xxxvn,  694-736,  tafeln 
xxvii-xxix. 

1886.  Die  Stegocephalen  aus  dem  Roth- 


liegenden des  Plauen'schen  Grundes  bei  Dresden. 
VI  Theil:  Die  Entwickelungsgeschichte  von 
Branchiosaurus  amblystomus  Cred.  Zeit.  d. 
deutsch.  geol.  Gesell.,  xxxvm,  576-633,  tafeln 
xvi-xix. 


188.  Credner,  Hermann.  1886.  Archegosaurus  Rcste 
aus  dem  Rothliegenden  v.  Offenbach.  Zeit.  d. 
deutsch.  geol.  Gesell.,  xxxvm,  696. 

1890.  Die  Stegocephalen  und  Saurier  aus 


189 


190. 


191. 


192. 


193- 


194. 


dem  Rothliegenden  des  Plauen'schen  Grundes  bei 
Dresden.  IX  Theil:  Hylonomus  geinitzi,  Petro- 
bates  truncatus,  Discosaurus  permianus.  Zeit. 
d.  deutsch.  geol.  Gesell.,  xlh,  240-277,  tafeln 
ix-xi,  figs. 

1887.     Stegocephalen  des  Rothliegenden. 


Wandtafeln  mit  Erlauterungen,  Leipzig.  Folio. 
Tafel  i,  Branchiosaurus  amblystomus;  Tafel  ii, 
Pelosaurus  laticeps  and  Melanerpeton  pulcher- 
rimum. 

.  1888.  Wandtafeln  mit  Stegocephalen  des 

Rothliegenden.  Neues  Jahrb.  f.  Mineral.  Geol. 
u.  Paleon.,  Bd.  I,  67-69. 

.  1 89 1.  Die  Urvierfussler  (Eotetrapoda)  des 

Sachsischen  Rothliegenden.  Allgemein-verstand- 
liche  naturwiss.  Abhandl.,  Heft  15.  Sonder- 
Abdruck  aus  der  "  Naturwisser.schaftlichen  Woch- 
enschrift,"  4-52,  53  figs. 

.  1893.  Die  Stegocephalen  und  Saurier  aus 

dem  Rothliegenden  des  Plauen'schen  Grundes  bei 
Dresden.  X  Theil:  Sclerocephalus  labyrinthicus 
H.  B.  Geinitz.  Zeit.  d.  deutsch.  geol.  Gesell., 
xlv,  639-704,  tafeln  xxx-xxxii. 

— - — .  1893.  Zur  Histologie  der  Faltenzahne 
l'aleozoischer  Stegocephalen.  Abhandl.  Sachs. 
Gesell.  Wiss.,  xx,  545-551. 

195.  .  1897.  Elemente  der  Geologic    Amphibia, 

489-494. 

196.  Cummins,  W.  F.  1908.  The  localities  and  horizons 

of  Permian  vertebrate  fossils  in  Texas.  Jour. 
Geol.,  xvi,  737-745- 

197.  Cuvier,    Georg.  1825.  Des   Batrachiens   fossiles. 

Recherches  sur  les  Ossemens  Fossiles,  3d  ed.,  431- 
444,  v,  pt.  u.     (Homo  diluvii  testis  et theoskopos.) 

198.  Davison,     Alvin.  1895.  A    contribution    to    the 

anatomy  and  phylogeny  of  Amphiuma  means. 
Jour.  Morphol.,  xi,  375-410,  2  pis.  (See  Kings- 
ley,  Tufts  College  Studies,  No.  7.) 

199.  Davis,    James   W.  1883.  On    the   occurrence    of 

remains  of  Labyrinthodonts  in  the  Yorkdale 
rocks  of  Wensleydale  (Carboniferous).  Rept. 
53d  Meeting  Brit.  Assn.  Adv.  Sci.,  Southport, 
Sept.,  492;  see  also  Nature,  xxvm,  578. 

200.  Dawson,  J.  W.  1853.  On  the  Coal  Measures  of  the 

South  Joggins,  Nova  Scotia.  Quart.  Jour.  Geol. 
Soc,  x,  1-41,  with  bibliography  of  papers  on  the 
coal  fields  of  Nova  Scotia. 

201.  •.  1855.  Acadian  geology.    Edinburgh,  1855, 

Amphibia,  160. 

.     1887.    Carboniferous   quadrupeds.     The 

Story  of  the  Earth  and  Man,  143-150.     9th  ed. 
1855.  Notice  of  the  discovery  of  a  reptilian 


202. 


203. 

skull  in   the   Coal   Measures  of  Pictou   (Nova 

Scotia).  Quart.  Jour.  Geol.  Soc,  xi,  8-9. 
204. .   1859.  New  species  of  reptiles  from  the  Coal 

Formation  of  Nova  Scotia.     Quart.  Jour.  Geol. 

Soc,  xvi,  273.     (Describes  Hylonomus.) 

i860.  On  the  terrestrial  Molluska,  a  chilog- 


205. 


206. 


nathus  Myriapod  and  some  new  species  from  the 
Coal  Formation  of  Nova  Scotia.  Quart.  Jour. 
Geol.  Soc,  xvi,  268. 

.  1862.  Additional  remains  of  land  animals 

in  the  Coal  Measures  of  Nova  Scotia.  Quart. 
Jour.  Geol.  Soc,  xvm,  5-7. 


BIBLIOGRAPHY   OF   THE   FOSSIL   AMPHIBIA. 


207 


2060. 


207. 


208.  - 


209. 


211. 
212. 

213- 

214.- 

2I.S- 
216. 

217. 

218. 

219. 


Dawson,  J.  W.  1862.  Descriptions  of  specimens  of 
fossil  Reptilia  discovered  in  the  Coal  Measures 
of  the  South  Joggins,  Nova  Scotia.  Quart.  Jour. 
Geol.  Soc.,  xvih,  238-244,  pis.  ix,  x. 

.  1863.  Note  on  the  footprint  of  a  reptile 

from  the  coal  of  Cape  Breton.  Canadian 
Naturalist  and  Geologist,  vm,  430-431,  1  fig.; 
review  in  Neues  Jahrb.  f.  Min.,  Geol.  u.  Paleon., 
64.  752- 

.  1863.  Air-breathers   of   the   Coal   Period. 

Montreal,  i-iv,  1-81,  pis.  1-6.  [A  descriptive 
account  of  the  remains  of  land  animals  found  in 
the  Coal  Formation  of  Nova  Scotia,  with  remarks 
on  their  bearing  on  the  theories  of  the  formation 
of  coal  and  of  the  origin  of  species.)  See 
also  Jour.  Geol.  Soc.,  xvi,  273;  xvm,  5-19, 
470;  also  Neues  Jahrb.  Mineral.,  Geol.  u.  Paleon., 
1864,  507. 

.  1 863.  Notice  of  a  new  species  of  Dendrerpe- 

ton  and  of  the  dermal  covering  of  certain  Carbo- 
niferous reptiles.  Quart.  Jour.  Geol.  Soc.,  xix, 
469-473- 

.  1863.  Air-breathers   of   the   Coal    Period. 

Canadian  Naturalist  and  Geologist,  vm,  1-12, 
81-92,  161-175,  268-295,  with  pis.  1-6. 

1 863.  The  Air-breathers  of  the  Coal  Period. 


Amer.  Jour.  Sci.  (2),  xxxvi,  430-432. 
— - — .   1868.  Land  animals  of  the  Coal   Period. 


Acadian  Geology,  1868,  2d  ed.,  chap,  xvm,  353- 
382. 

1870.  Note  on  some  new  animal  remains 


from  the  Carboniferous  and  Devonian  of  Canada. 
Geol.  Mag.,  vil,  87;  Quart.  Jour.  Geol.  Soc., 
xxvi,  166-167;  Canadian  Naturalist  (2),  v,  98-99. 
(Same  paper  in  all:  Baphetes  minor.) 
.  1876.  On  a  recent  discovery  of  Carbonifer- 
ous Batrachians  in  Nova  Scotia.  Amer.  Jour.  Sci. 
(3),  xii,  440-447- 

1878.  Acadian  geology.    Supplement  to  3d 


223. 


edition,  London,  Amphibia,  57-65. 
.  1882.  On  the  results  of  the  recent  explora- 
tions of  the  erect  trees  containing  animal  remains 
in  the  Coal  Formation  of  Nova  Scotia.  Phil. 
Trans.  Roy.  Soc.  London,  clxxiii,  pt.  11,  621- 

654.  Pis-  39-47- 

.   1882.  On  the  results  of  recent  explorations 

of  the  erect  trees  (Batrachia).  Canadian  Natu- 
ralist (2),  x,  252-254.    (Abstract.) 

.   1 89 1.  On  new  specimens  of  Dendrerpeton 

acadianum,  with  remarks  on  other  Carboniferous 
Amphibians.  Geol.  Mag.  (3),  vm,  145-156, 
4  figs. 

.  1891.  Note    on    Hylonomous    lyelli,  with 

photographic  reproduction  of  skeleton.  Geol. 
Mag.,  n.  s.,  Dec,  m,  vm,  258-259,  pi.  8. 

.   1 89 1.  On  the  mode  of  occurrence  of  remains 

of  land  animals  in  erect  trees  of  Nova  Scotia. 
Trans.  Roy.  Soc.  Canada,  ix.  sec.  IV,  127-128. 

.   1894.  Some  salient  points  in  the  science  of 

the  earth.  The  Oldest  Air-breathers,  chap,  x, 
259-307,  with  restorations  and  figures. 

.  1894.  Preliminary  note  on  the  recent  dis- 
coveries of  Batrachians  and  other  air-breathers  in 
the  Coal  Formation  of  Nova  Scotia.  Canadian 
Record  of  Science,  vi,  1-7,  I  fig. 

.  1895.  Synopsis  of  the  air-breathing  ani- 
mals of  the  Paleozoic  of  Canada  (up  to  1894). 
Trans.  Roy.  Soc.  Canada,  XII,  sec.  IV,  71-78. 


224. 
225. 

226. 

I   227. 
228. 

229. 

230. 

231- 
232. 

233- 
234- 
235- 
236. 

237- 
238. 

239- 
240. 


Deichmieli.er,  J.  V.  1882.  Ueber  "Die  Stego- 
cephalen  aus  dem  Rothliegenden  des  Plauen'schcn 
Grundes  bei  Dresden.  Ill  Theil:  von  H.  Cred- 
ner."    Sitzungsb.  Isis,  Dresden,  71. 

.  1884.  Ueber      Branchiosaurus      petrolei, 

Nachtrage  zur  Dyas  III.  Mittheilungen  des  kgl. 
mineral,  geol.  u.  praehist.  Museum  in  Dresden, 
Heft  6,  Paleontol.  Abhandl.  aus  dem  Dresdener 
Museum,  5-17,  Taf.  1. 

Dollo,  Louis.  1884.  Note  sur  le  Batracien  de 
Bernissart.  Bull.  Mus.  Roy.  Hist.  Nat.  Belg  , 
m.  85~93.  pl-  i»-  (Hylasobatrachus  croyii  from 
the  Wealden.) 

.  1885.  Les  Labyrinthodontes.    Rev.  Quest. 

Scientif.  Bruxelles,  xvn,  305-312. 

■  '895.  Sur  la  Phylogenie  des  Dipneustes. 

Bull.  Soc.  Beige  de  Geol.  Paleont.  Hydrol.,  ix, 
79-178,  pis.  v-x.  (Discusses  relations  to  the 
Amphibia.) 

Dunlop,  Robert.  1910.  The  fossil  Amphibia  in  the 
Kilmarnock  Museum  previous  to  the  fire  of 
November  26,  1909.  Annals  Kilmarnock  Glen- 
field  Ramblers  Society,  No.  6,  10,  2  pis. 

Eastman,  Chas.  R.  1902.  The  Carboniferous  fish 
fauna  of  Mazon  Creek,  Illinois.  Jour.  Geol.,  x, 
535-541.  with  list  of  fossil  vertebrates. 

.  1902.  Translation  of  Zittel's  Text-book  of 

Paleontology,  vol.  II,  114-139,  figs.  193-235. 

-.  1907.  Devonic   fishes   of   the   New   York 


Formations.     Memoir  No.  10,  New  York  State 

Mus.,  180-183. 
Eichwald,  E.  1848.  On  Zygosaurus  lucius  from  the 

Permian  of  Russia.     Bull,  de  la  Soc.  imp.  des 

Natl,  de  Moscou,  xxi,  159. 
.  i860.  Letha?a  rossica  ou  Paleontologie  de 

la  Russie,  I,  Stuttgart.    1620-1633,  tafeln  57-59, 

Labyrinthodonten. 
Embleton  and  Atthey.   1874.  On  the  skull  and 

other  bones  of  Loxomma.    Ann.  and  Mag.  Nat. 

Hist.,  ser.  4,  xiv,  38-63,  pis.  iv-vii. 
Emery,  C.  1897.  Die  fossilen  Reste  von  Archego- 

saurus  und  Eryops  und  ihre  Bedeutung  fur  die 

Morphologie  des  Gliedmassenskelets.   Anat.  Anz., 

xiv,  No.  8,  201-208,  figs.  1-7. 
Emmons,  E.   1857.  Perm  und  Trias  System  in  Nord 

Carolina.    Edinb.  n.  Phil.  Jour.,  v,  370. 
.  1857.  Dictyocephalus       elegans       Leidy. 

American  Geology,  pt.  vi,  Labyrinthodonts,  59, 

figs-  31-32- 

-.  1863.  Dictyocephalus  elegans  Leidy.  Man- 


242. 


243- 


ual  of  Geology,   New  York.     Labyrinthodonts, 

180,  184. 
Emmrich,    A.    1852.   Bericht  ueber  die  Naturfor- 

scherversammlung  in  Gotha  im  Jahre  1851.  Jahrb. 

d.  k.  k.  geol.  Reichsanstalt,  p.  155,  Wien. 
Etheridge,   Robert.  1866.  On   the   discovery  of 

several  new  Labyrinthodont  reptiles  in  the  Coal 

Measures  of  Ireland.     Geol.  Mag.,  m,  4.     (See 

Huxley,  1867.) 
Fraas,  E.  1S89.  Die  Labyrinthodonten  der  Schwtb- 

iscben   Trias.     Paleontographica,   XXXVI,    1-158, 

with  bibliography. 
.   1896.  Die     Schwabischen     Trias-Saurier. 

Fcstgabe    des    koniglichen   naturalien  Cabinets 

in  Stuttgart  zur  42  Versammlung  dcr  deutsch. 

geol.  Gcscll.,  Stuttgart,  1-8,  pis.  i-ii. 


208 


THE   COAL   MEASURES  AMPHIBIA   OF   NORTH   AMERICA. 


244.  Fraas.E.  1901.  Labyrinthodon  aus  dem  Buntsand- 
stein  von  Teinach.  Jahreshefte  des  Vereins  f. 
vaterlandische  Naturkunde  in  Wurtemberg,  Jahr., 

LVH,  318-320. 

-.  1903.  Die  geognostische  Sammlung  Wur- 


245- 


246. 


247. 


tembergs  im  Parterre-Saal  zugleich  ein  Leitfaden 
fur  die  geologischen  Verhaltnisse  und  die  vor- 
weltlichen  Bewohner  unseres  Landes.  2d  Auflage. 
1903.  Rana  danubina  H.  von  Meyer  var. 


rara,  0.  Fraas  aus  dem  Obermiocan  von  Stcin- 
heim.  Jahreshefte  des  Vereins  f.  vaterl.  Natur- 
kunde in  Wurtemberg,  Jahr.  1903,  105-110, 
with  fig. 

-.  1909.  Rana  haufiftana  n.  sp.  aus  den  Dyso- 


dilschiefern  des  Randecker  Maares.  Jahreshefte 
des  Vereins  f.  vaterl.  Naturk.  in  Wurtemberg, 
Jahr.  1909,  7,  with  1  fig. 

247a. .  1913.  Neue    Labyrinthodonten    aus    der 

schwabischen  Trias.  Paleontographica,  Bd.  lx, 
275-294,  pis.  xvi-xxii,  with  text-figs.  1-5. 

248.  Frech,     Fritz.  1901.  Lethaea     geognostica     oder 

Beschreibung  und  Abbildung  der  fur  die  Gebirgs- 
formation  bezeichnendsten  Versteinerungen.  I 
Theil,  Lethaea  palaaozoica,  2  band.  Die  Dyas, 
459-471.  22  figs.,  pi.  56. 

249.  Fritsch,    Anton.  1870.  Sitzungsberichte    der    k. 

Gesell.  d.  Wissen.,  April  27,  1870.  (Describes  2 
new  species  of  Amphibia.) 

1875.  Ueber  die  Fauna  des   Pilsner  und 


250. 


251 


252. 


253- 


Rakonitzer   Beckens.    Sitzungsb.   d.   k.  Boehm- 
ischen  Gesell.  d.  Wissenschft.,  March. 

1883-1899.  Fauna  der  Gaskohle  und  der 


Kalksteine  der  Permformation  Boehmens.  Bd.  1, 
1-182,  116  figs.  Tafeln  i-xlviii;  Bd.  II,  1-64,  figs. 
1 17-142,  Tafeln  xlix-lxx;  Bd.  IV,  supplement  85- 
98,  figs.  383-394,  Tafeln  clxiii-clxiv.  This  is  the 
chief  work  on  extinct  Amphibia. 

189,5.  Ueber   neue    Wirbelthiere    aus    der 


Permformation  Boehmens,  nebst  einer  Ueber- 
sicht  der  aus  derselben  bekannt  gewordenen 
Arten.  Sitzungsber.  d.  k.  Boehmisch.  Gesell.  der 
Wissenschaften,  Prag,  1. 

-.  1905.  Vorlaufige  Notiz  ueber  .Miscellanea 


Paleontologica  aus  Boehmen  und  America. 
Sitzungsber.  d.  k.  Boehm.  Gesell.  dej  Wissen- 
schaften, Prag,  Kammplatten,  3. 

254.  Fritsch,    Karl    Freiherr.    1879.  Referat    ueber 

Gaudry's  Sur  les  Reptiles  des  Temps  primier; 
Nachweis  von  Protriton  petrolei  bei  Oberhof  in 
Thueringen.  Neues  Jahrb.  f.  Mineral.  Geol.  u. 
Paleon.,  720. 

255.  Fuchs,     Hugo.     1909.     Betrachtungen  ueber  die 

Schlafengegend  am  Schadel  der  Quadrupeda. 
Anat.  Anz.,  xxxv,  No.  5/7,  1 13-167,  25  figs. 

256.  Gadow,    Hans.  1896.  On    the    evolution    of    the 

vertebral  column  of  Amphibia  and  Amnipta. 
Phil.  Trans.  Roy.  Soc.  London,  ser.  B,  clxxxvii, 
I_57»  5°  figs-.  and  bibliography. 

257.  .  1901.  Amphibia  and  reptiles.     Cambridge 

Nat.  Hist.,  vm,  Stegocephala,  80.    London. 

258.  Gaudry,  Albert.  1866.  Note  sur  le  Reptile  dS- 

couvert  a  Muse.  Comptes  Rendus  de  l'Academie 
de  Science,  August,  341. 

259.  .  1867.  M6moire  sur  le  Reptile  decouvert 

par  M.  Frossard  a  Muse.  Nouv.  Archiv  du 
Musee  d'Hist.  Nat.,  m,  22-40,  pi.  iii. 

260.  -       — .    1868.    Note    sur    L'Actinodon    frossardi. 


261. 
262. 

263. 

264. 

265. 

266. 

267. 
268. 

269. 

270. 

271. 
272. 
273- 

274. 

275- 
276. 

277- 

278. 
279. 

280. 

281. 
282. 

283. 


Gaudry,  Albert.  1874.  Les  Ettcs  de  Temps  prim- 
aire.  Revue  Scientifique,  ser.  2,  XIII,  993. 

.  1875.  Les   Reptiles   des   Schistes    bitumi- 

neaux  dAutun.  Bull.  Soc.  Geol.  de  France,  3d 
ser.,  iv,  720. 

-.  1875.  Sur   la    Decouverte    de   Batraciens 

proprement  dits  dans  le  Terrain  primaire.  Comp- 
tes Rendus,  lxxx,  441-443. 

.  1875.  Sur   la    Decouverte    de    Batraciens 

dans  le  Terrain  primaire.  Bull,  de  la  Soc.  Geol. 
de  France,  ser.  3,  m,  300-306,  pis.  vii-viii.  (Sec 
also  263  above;  Jour.  Zool.,  iv,  38-41,  342.) 

.     1876.     Les  Reptiles  des  Schistes  bitumi- 

neux  d'Autun.  Bull.  Soc.  Geol.  de  France,  ser. 
Ill,  IV,  720-723,  pi.  xxii;  1879,  VII,  62;  1885,  3d 
ser.,  xm,  44. 

.  1878.  Sur  les  Reptiles  de  Temps  primaires. 

Comptes  Rendus,  lxxxvii,  956. 

1878.   Les  Vertebres  fossiles  des  environs 


dAutun.    Bull.  Soc.  Geol.  France,  vn,  62. 

-.  1878.  Les  Reptiles  d.  l'Epoque  Permienne 


aux  Environs  d'Autun.  Bull.  Soc.  Geol.  de  France, 
ser.  3,  vii,  62-76,  pis.  iii-iv. 

.  1880.   Sur    un    Reptile    tres    perfectionne' 

trouve'    dans    le    terraine    Permien.      Comptes 
Rendus,  xci,  669-671. 

— .  1 88 1.  Sur  les  plus  anciens  Reptiles  trouves 


en  France.     Comptes  Rendus  Acad.  Sci.  Paris 
xcn,  1143-1145. 

.  1 88 1.  On  a  highly  organized  reptile  from 

the  Permian  Formation.    Amer.  Nat.,  vn,  67-71. 
1883.  Les  Reptiles  primaires.    Arch.  Zool. 


Exper.,  2d  ser.,  I,  5-30,  pis.  1-7.  with  figs. 

1 883.  Les  Enchainements  du  Monde  animal 


dans  les  Temps  geblogiques.    Fossiles  Primaires, 

with  4  figs.     Paris. 
.  1884.  Nouvelle     Note    sur    !es    Reptiles 

Permiens.     Bull.  Soc.  Geol.  de  France,  3  ser., 

xm,  44-51,  pis.  iv-v.    Paris. 
.  1884.  Nouvelle     Note    sur    les    Reptiles 

Permiens.      Comptes    Rendus  Acad.  Sci.  Paris, 

xcix,  737-738. 
— .  1885.  Paleontology     in     Germany     and 


America.    Revue  Sci.,  7,  Paris;  Geol.  Mag.  (3), 
11,  London. 

1886.  Sur  un  nouveau  Genre  de  Reptile  du 


Permien  d'Autun.  Bull.  Soc.  Geol.  de  France,  ser. 
in,  xiv,  430-433,  taf.  23. 
.  1887.  L'Actinodon.      Nouvelles    Archives 


Mus.  Hist.  Nat.,  Paris,  2d  ser.,  x,  30  pp.,  3  pis. 
.  1888.  Les  Vertebres  fossiles  des  environs 

d'Autun.     Mem.  Soc.  d'Hist.  nat.  d'Autun,  90 

pp.  et  6  planches. 
.  1885.  Sur    les    Similitudes    que    plusiers 

Reptiles  ont  eues  dans  divers  Pays  de  Monde  ver 

la  Fin  des  Temps  primaires.    Compte  Rendus  de 

la  3d  session  Congres  gcologique  international  a 

Berlin,  1-4. 
.    1888.    Les  Vertebras  fossiles  des  environs 

d'Autun.     Autun. 

1 890.  Les  Enchainement  du  Monde  Animal 


Bull.  Soc.  Geol.  de  France,  Ser.  2,  xxv,  576. 


dans  les  Temps  Geologiques,  Fossiles  Secondaires. 
Amphibia,  169. 
Gaupp,  Ernst.  1895.  Beitrage  zur  Morphologie 
des  Schadels.  Ill:  Zur  vergleichenden  Anatomie 
der  Schlafengegend  am  knochernen  Wirbelthier- 
Schadel.  Morphologische  Arbeiten  herausge- 
geben  von  G.  Schwalbe,  iv,  1,  77-128. 


BIBLIOGRAPHY   OF  THE  FOSSIL  AMPHIBIA. 


209 


284.  Gegenbaur,  Carl.  1862.  Untersuchungen  zur  ver- 

gleichen.  Anatomic  dcr  Wirbelsaule  bei  Amphi- 
bien  und  Reptilien.     Leipzig. 

285.  .  1895.  Clavicula  und  Cleithrum.    Morphol. 

Jahrb.,  xxm,  1-20. 

286.  Geikie,    ARCHinALD.  1905.  Scheuchzer's    "Fossil 

Man."  Founders  of  Geology,  2d  ed.,  98-99. 
London. 

287.  Geinitz,  H.  B.  1864.  Palaeosiren    beincrtii   Gein. 

ein  neues  Reptile  aus  der  unteren  Dyas  von 
Oelberg  bei  Braman.  Neues  Jahrb.  f.  Mineral., 
Geol.  u.  Paleon.,  513-516. 

288.  .  1885.  Ueber  Thierfahrten  in  der  Steinkohl- 

enformation  von  Zwickau.  Festschrift  Natur. 
Gesell.  Isis,  63-66,  pi.  2. 

und  Deichmueller.  1882.  Ueber  die  fos- 


289. 


290. 


silen  Saurier  in  dem  Kalke  des  Rothliegenden  bei 
Dresden.     Sitzungsber.  Isis,  6-9. 

1882.  Die  Saurier  des  unterer  Dyas  von 


Sachsen.    Paleontographica,  xxix,  9-46,  pis.  1-9. 

291.  Gergens,  Dr.  1844.  Letter    to    Professor    Bronn 

announcing  the  discovery  of  the  Apatheon  de- 
scribed by  von  Meyer  as  an  amphibian.  The 
first  announcement  of  Amphibia  from  Carbonifer- 
ous. Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  49. 

292.  Giebel,  C.  G.  1847.  Fauna  der  Vorwelt  mit  steter 

Berucksichtigung  der  lcbenden  Thiere.  Am- 
phibia in  Bd.  I,  Abth.  2,  165-213. 

293.  .  1854.  Odontography,  Leipsic. 

294.  Goldenburg,  Fr.  1873.  Fauna  Saraepontana  fos- 

silis — Die  fossilen  Thiere  aus  der  Steinkohlenfor- 
mation  von  Saarbrucken.  Heft  I,  p.  4,  Taf.  1, 
fig.  1.     (Anthracosaurus  raniceps.) 

295.  Goldfuss,  Dr.  A.  1831.  Beitrage   zur  Kenntniss 

verschiedener  Reptilien  der  Vorwelt.  Reptilien 
aus  der  schieferigen  Braunkohle.  Verhandlungen 
der  k.  leop.  Carol.  Akad.  d.  Naturf.,  xv,  Abth.  1, 
1 17-126,  pis.  xii-xiii. 

-.  1847.  Ueber  das  alteste  der  mit  Bestimmt- 


296. 


297. 


heit  erkannten  Reptilien .  Neues  Jahrb.  f .  Mineral. , 
Geol.  u.  Paleon.,  400,  taf.  6.    (Archegosaurus.) 
1847.  Beitrage   zur    vorweltlichen    Fauna 


des  Steinkohlengebirgen.    Bonn,  quarto,  taf.  1-3. 

298.  Goodrich,  E.  S.  191  i.  On  the  segmentation  of  the 

occipital  region  of  the  head  in   the  Batrachia 
Urodela.    Proc.  Zool.  Soc.  London,  101-120,  figs. 

299.  Gregory,  W.  K.  191 1.  The  limbs  of  Eryops  and 

the  origin  of  paired  limbs  from  fins.     Science, 
n.s.,  xxxin,  No.  848,  508-509. 

1913.  Crossopterygian     ancestry    of    the 


300. 


3000. 


Amphibia.    Science,  n.  s.,  xxxvn,  No.  960,  806- 
808. 

-.  1915.  Present  status  of  the  problem  of 


the  origin  of  the  Tetrapoda,  with  special  reference 
to  the  skull  and  paired  limbs.  Ann.  New  York 
Academy  of  Science,  XXVI,  3I/-383.  P'-  iv. 
(Stegocephali,  p.  365.) 

301.  Gunther,    Albert.  1889.  On    sexual    differences 

found  in  bones  of  some  recent  and  fossil  species 
of  frogs  and  fishes.  Ann.  and  Mag.  Nat.  Hist., 
ser.  3,  iv,  377. 

302.  Gurich,  G.  1884.  Ueber  einige  Saurier  des  ober- 

schles.  Muschelkalkes.  Zeit.  d.  deutsch.  geol. 
Gesell.,  xxxvi,  141. 

303.  Hancock,  A.  1869.  Note  on  Anthracosaurus.   Ann. 

and  Mag.  Nat.  Hist.,  4th  ser.,  iv,  270. 


304.  Hancock  and  Atthey.  1868.  Notes  on  the  remains 
of  some  reptiles  and  fishes  from  the  shales  of  the 
Northumberland  Coal  Field.  Ann.  and  Mag. 
Nat.  Hist.,  4th  Ser.,  I,  266-278,  pis.  xiv-xvi. 

3°5-  •  1869.  On  a  new  Labyrinthodont  Amphib- 
ian from  the  Northumberland  Coal  Field  and  on 
the  occurrence  in  the  same  locality  of  Anthraco- 
saurus. Ann.  and  Mag.  Nat.  Hist.,  ser.  4,  iv, 
182-189;  also  Nat.  Hist.  Trans.  Northumberland 
and  Durham,  ill,  1869,  330.  (Urocordyhts 
reticulatus.) 

.  1870.  On   Urocordylus,    Pteroplax.     Nat. 

Hist.  Trans.  Northumberland  and  Durham,  in, 


306. 
307. 
308. 

309. 

310. 
3"- 

312- 

313. 
314 

315- 
316. 

317- 

318. 
319- 
320. 

321. 

322. 


30.  79- 

.  1870.  On  the  occurrence  of  Loxomma  all- 

manni  in  the  Northumberland  Coal  Field.  Ann. 
and  Mag.  Nat.  Hist.,  ser.  4,  v,  374-379. 

.  1870.  Description    of    a    Labyrinthodont 

Amphibian;  a  new  generic  form  obtained  in  the 
Coal  Shales  at  Newsham.  Ann.  and  Mag.  Nat. 
Hist.,  ser.  iv,  vi,  56-65,  pi.  i.  (Batrachiderpeton 
lineatum.) 

.  1 87 1.  Description  of  a  considerable  por- 
tion of  a  mandibular  ramus  of  Anthracosaurus 
russelli,  with  notes  on  Loxomma  and  Archichthys. 
Ann.  and  Mag.  Nat.  Hist.,  4th  ser.,  vn,  73,  pi.  vi. 

.  1871.  Batrachiderpeton.    Nat.  Hist.  Trans. 

Northumberland  and  Durham,  rv,  208. 

Hancock  and  Howse.  1870.  On  a  new  Labyrintho- 
dont Amphibian  from  the  magnesium  limestone 
of  Midderidge,  Durham.  Quart.  Jonr.  Geol. 
Soc.,  xxvi,  556-564,  pi.  38.  Permian  of  Eng- 
land, Lepidotosaurus  diiffii,  a  Labyrinthodont  of 
some  3  to  4  feet  in  length. 

Haeckel,  Ernst.  1895.  Systematische  Phylogenie 
der  Wirbelthiere  (Vertebrata).  Amphibia,  266- 
283.  Also  History  of  Creation,  Trans.,  5th  ed., 
1909,  11,  295-300. 

Hatch  and  Corstorphine.  1905.  The  geology  of 
South  Africa.    The  Karoo  System.  194. 

Hatcher,  J.  B.  1905.  The  vertebrate  fauna  of  the 
Judith  River  Beds.  Bull.  U.  S.  Geol.  Surv.,  No. 
257,  67-103.     (Scapherpeton  and  Hemitrypus.) 

Hay,  O.  P.  1890.  The  skeletal  anatomy  of  Amphi- 
uma  during  its  early  stages.  Jour.  Morphol.,  iv, 
1  i_34i  with  pis. 

.  1900.  Description  of  some  vertebrates  of 

Carboniferous  Age.  Proc.  Amer.  Phil.  Soc., 
xxxix,  No.  161,  96-123.  (Amphibamus  grandi- 
ceps.) 

.  1902.  Catalogue  and  bibliography  of  fossil 

Vertebrata  of  North  America.  Bull.  U.  S.  Geol. 
Surv.,  No.  179,  Batrachia,  409-425. 

Hickling,  G.  1909.  British  Permian  footprints. 
Mem.  and  Proc.  Manchester  Lit.  and  Phil.  Soc, 

53.  3°.  Pi-  3- 
Hilton,  Wm.  A.  1902.  A  structural  feature  con- 
nected with  the  mating  of  Diemyctylus  virides- 

cens.    Amer.  Nat.,  xxxvi,  643. 
Hoepen,   E.  C.  N.  van.  1915.  Stegoccphalia   of 

Senekal.    Ann.  of  the  Transvaal  Mus.,  v,  No.  2, 

125-149,  pis.  xvi-xxiv. 
Hoernes,  Rud.  1884.  Elcmente  der  Paleontologic, 

445-453-    Leipzig. 
Hoffman,  A.  H.  1873-78.  Amphibien.    Bd.  Vi,  2d 

Abth.,    Bronn's    Classen    und    Ordnungen    des 

Thierreiches. 


210 


THE    COAL    MEASURES   AMPHIBIA   OF   NORTH    AMERICA. 


323.  Huene,   Friedrich   von.  1902.  Uebersicht   ueber  i 
die  Reptilien   der  Trias.     Geol.  u.    Paleon.  Ab- 
handl.,  N.  F.,  vi,  H.  i,  20,  21 ,  81. 

igio.    Neubeschreibung     des     Permischen 


324 


325- 
326. 


327- 


Stegocephalen-Dasyceps  bucklandi  aus  Kenil- 
worth.  Geol.  u.  Paleon.  Abhandl.  herausgegeben 
v.  E.  Koken,  N.  F.,  vm,  H.  6,  33-46,  fig.  pi.  i,  ii. 

.  1912.  Beitrage  zur  Kenntniss  des  Schadels 

von  Eryops.    Anat.  Anz.,  Bd.  41,  pp.  98-104,  figs. 
-.  1913.  Ueber   Lysorophus   aus   dem    Perm 


von  Texas.    Anat.  Anz.,  Bd.  43,  389-396. 
— .   19 13.  The  skull  elements  of  the  Permian 


Tetrapoda  in  the  American  Museum  of  Natural 
History,  New  York.  Bull.  Am.  Mus.  Nat.  Hist., 
xxxii,  315-386,  57  figs. 
328.  Huxley,  Thomas  Henry.  1859.  On  Dasyceps 
bucklandi  (Labyrinthodon  bucklandi  Lloyd). 
Mem.  Geol.  Surv.  United  Kingdom,  52-56,  figs. 
1-2;  Scientific  Memoirs,  11,  263,  figs.  1-2. 

1859.  On  a  fragment  of  a  lower  jaw  of  a 


329- 


33°- 


331- 


332. 


333- 


334- 


335- 


336. 


large  Labyrinthodont  from  Cubbington.  Mem. 
Geol.  Surv.  United  Kingdom,  56-57;  Scientific 
Memoirs,  11,  269. 

1859.  On  some  Amphibian  and  reptilian 


remains  from  South  Africa  and  Australia.    Quart. 
Jour.    Geol.    Soc.,    xv,    642-649,   pis.   xxi-xxii; 
Scientific  Memoirs,  11,  120-129,  p's.  7-8. 
.  1862.  On  new  Labvrinthodonts  from  the 


Edinburgh  Coal  Field.  Quart.  Jour.  Geol.  Soc., 
xvm,     291-296,   pi.   xi;   Scientific   Memoirs,   11, 

53""535.  Pi-  37- 
.  1863.  Description  of  Anthracosaurus  rus- 

selli,  a  new  Labyrinthodont  from  the  Lanark- 
shire Coal-Field.  Quart.  Jour.  Geol.  Soc,  xix, 
56-68,  figs.;  Scientific  Memoirs,  11,  558-572,  figs. 
1-2. 

.  1865.  On  a  collection  of  vertebrate  fossils 

from  the  Panchet  Rocks,  Ranigunj.  Bengal. 
Mem.  Geol.  Surv.  of  India;  Paleontologia  Indica, 
ser.  iv;  Indian  Pretertiary  Vertebrata,  t,  Am- 
phibia. 3-8,  pis.  vi,  and  figs.;  Scientific  Memoirs, 
Hi,  90-120,  figs.,  pi.  16. 

1867.  On  a  collection  of   fossil  Vertebrata 


337- 


from  the  Jarrow  Colliery,  County  of  Kilkenny, 
Ireland.  Trans.  Roy.  Irish  Acad.,  1871,  xxiv, 
351-369  (Read  Jan.  6,  1866),  pis.  xix-xxiii;  Scien- 
tific Memoirs,  in,  180-197,  pis.  17-21. 

.  1869.  On    a    new    Labyrinthodont    from 

Bradford.  Quart.  Jour.  Geol.  Soc.  London,  xxv, 
309-311,  pi.  xi;  Scientific  Memoirs,  III,  391-393, 
pi.  26. 

.  1875.   Amphibia.   Art.   in    "Encyclopedia 

Britannica,"  9th  ed.,  I,  750-771. 

.   1879.    On  the  characters  of   the  pelvis  in 

the  Mammalia  and  the  conclusions  respecting  the 
origin  of  mammals  which  may  be  based  on  them. 
Proc.  Roy.  Soc.  Lond.,  xxvm,  395-405,  figs. ; 
Scientific  Memoirs,  iv,  345-356,  figs. 
338.  Jaeger,  Dr.  Georg  Friedrick.  1824.  De  Ichthyo- 
sauri sive  Proteosauri  Fossilis  Specimenibus  in 
Agro  Bollensi  in  Wurtembergia.  Folio,  11.  (First 
mention  of  the  discovery  of  Labyrinthodonts,  a 
tooth  and  occipital  region  of  a  cranium,  which  in 
1 833  Jaeger  designated  Mastodonsaurus  giganteus. 
This  was  and  still  is  the  largest  known  amphibian.) 

339-  — ; •  1828.  Ueber  die  fossilen  Reptilien  welche 

im  Wurtemburg  aufgefunden  worden  sind. 
Quarto,  35-38,  pis.  iv-v. 


340.  Jaeger,  Dr.  G.  F.  1833.  Mastodonsaurus  gigan- 
teus.    Bull,  de  la  Soc.  Geol.  de  France,  HI,  86. 

341. .   1850.  Ueber    die    Uebereinstimmung    des 

Pygopterus  lucius  Agassiz  mit  dem  Arehego- 
saurus  decheni.  Abh.  Math-phys.  Classe  d.  Bayr. 
Akad.,  v,  877,  pi. 

342.  Jaekel,  Otto.  1894.  Ueber  die  sogennante  Falten- 

zahne  und  complicirte  Zahnbildung  ueberhaupt. 
Sitzb.  d.  Gesell.  naturf.  Freunde  zu  Berlin,  No.  5, 
146. 

343.  .  1896.  Ueber   die    Korperform   und    Haut 

Bedeckung  von  Stegocephalen.  Sitzungsber.  d. 
Gesell.  naturf.  Freunde  zu  Berlin,  1-8,  1  fig. 

344. .   1896.  Die  Organization  von  Archegosaurus. 

Zeit.  d.  deutsch.  geol.  Gesell.,  xlviii,  505-521, 

figs.  1-10. 
345. .   1902.  Gephyrostegus  bohemicus.      Zeit.  d. 


346. 


347- 


348. 


349- 


35o. 


351- 


deutsch.  geol.  Gesell.,  liv,  Heft  3,  127-132. 
.   1903.  Ueber  die  Epiphyse  und  Hypophyse. 

Sitzungsb.    d.    Gesell.    naturf.    Freunde,    No.    2, 

27-58. 
.   1903.  Ueber  Ceraterpeton,  Diceratosaurus 

und  Diplocaulus.  Neues  Jahrb.  f.  Mineral.,  Geol. 

u.  Paleon.,  Bd.  I,  109-134,  pis.  ii-v. 
.   1904.  Ueber  die  Bildung  der  ersten  Hals- 

wirbel    und    die  Wirbelbildung  im  Allgemeinen. 

Zeit.  d.  deutsch.  geol.  Gesell.,  lvi,  109-119,  figs. 
.   1905.  Ueber  die   primare   Gliederung  des 

Unterkiefers.      Sitzungsber.    d.    Gesell.    naturf. 

Freunde,  No.  4,  134. 
.   1909.  Ueber  die  Klassen  der  Tetrapoden. 

Zool.  Anz.,  xxxiv,  Nr.   7/8,  Apr.  20,   193-212, 

figs.  1-15. 

1909.  Ueber  die  Beurtheilung  der  paarigen 


352- 


Extremitaten.  Sitzungsb.  d.  konig.  Preuss.  Akad. 
d.  Wissen.,  xxvi,  707-724. 

.   1913.  Ueber  den  Bau  des  Schaedels.    Anat. 

Anz..  Erganzungsheft,  Bd.  44,  77-94.  mit  figuren. 

353.  Jordan.  H.  1849.  Erganzende  Bemerkungen  zu  der 

Abhandlung  von  Goldfuss  ueber  die  Gattung 
Archegosaurus.  Verhandl.  d.  naturhist.  Vereines 
f.  Rheinlande  u.  Westphalen,  vi,  76-81,  taf.  4. 

354.  Kayser,  Friedrich.  1902.  Geologie  von  Boehmen. 

Verlag  von  I.  Taussig,  1074-1210.  (Formations 
of  Bohemia  where  Amphibia  have  been  found.) 

355.  Kinahan,  G.  H.   1878.  Geology  of  Ireland,  London; 

The  Jarrow  Colliery.     116-119. 

356.  King,  Alired  T.   1844.  Description  of  fossil  foot- 

marks supposed  to  be  referable  to  the  classes 
Birds,  Reptilia.  and  Mammalia  found  in  the 
Carboniferous  series  in  Westmoreland  County, 
Pennsylvania.      Proc.    Acad.    Nat.    Sci.,    Phila., 

175-179- 

357.  .   1845.  Description  of  a  fossil  footmark  of 

the  quadruped  Thenaropus.  Amer.  Jour.  Sci., 
xlviii,  343-352,  12  figs.;  also  1  (n.  s.),  268. 

358.  Kingsley,  J.  S.  Systematic  position  of  Coecilians. 

Tufts  College  Studies  No.  7  (xxx);  also  No.  7 
(xxvm),  Amphiuma. 

359.  Ki.aatsch,    Hermann.    1896.  Die   Brustflosse  der 

Crossopterygier.  Festschrift  fur  Gegenbaur,  Bd. 
1,  259-391,  Taf.  i-iv. 

360.  Knipe,  Henry  R.  1912.  Amphibians  of  the  Car- 

boniferous.   Evolution  in  the  Past,  59,  and  plate. 

361.  Kunisch,  Hermann.  1885.  Ueber  die  Unterkiefer  von 

Mastodonsaurus  silesiacus  Hun.  Zeit.  d.  deutsch. 
geol.  Gesell.,  xxxvn,  528-533,  with  figs.  1-2. 


BIBLIOGRAPHY  OF   THE  FOSSIL   AMPHIBIA. 


211 


362.  Kunisch,    Hermann.     1890.     Labyrimhodonten, 

Reste  d.  oberschles.  Musehelkalkes.  Zeit  d. 
deutsch.  geol.  GcmD.,  xi.ii,  377-385.  Taf.  xx. 

363.  Kutorga,  Stefh.  1838.  Bcitrag  zur  Kenntniss  der 

organ.  Ueberreste  des  Kupfersandsteines  am  west- 
lichen  Abhange  des  Urals.  St.  Petersburg,  rait 
tafeln. 

364.  Lake,  J.  J.  1878.  Reptiles  of  the  primary  period. 

English  Mechanic,  xxvm,  438-439. 

365.  Lambe,     L.     M.   1902.  Scapherpeton     tectum — a 

batrachian  from  the  Cretaceous.  Contr.  Cana- 
dian Paleontology,  ill,  pt.  11,  31. 

366.  .   1904.  Progress  of  vertebrate  paleontology 

in  Canada.    Trans.  Roy.  Soc.  Canada,  x,  sec.  iv, 

I3-56- 

367.  Laube,    Gustav    C.  1898.  Andriasreste    aus    der 

Boehmischen  Braunkohlenformation.  Abhandl. 
d.  deutsch.  naturw.-med.  Vereines  f.  Boehmen 
"Lotos,"  1,  heft  2,  25. 

1898.  Amphibienreste  aus  dem  Diatoma- 


368. 


369- 


ceenschiefer  von  Sulloditz.  Abhandl.  d.  deutsch. 
naturw.-med.  Vereines  f.  Boehmen  "Lotos,"  1, 
heft  3,  10,  taf.  i,  fig.  6. 

1 901.  Synopsis  der  Wirbelthierfauna  der 


bohm-Braunlcohlenformation  und  Beschreibung 
neuer  oder  bisher  unvollstandig  bekannter  Arten. 
Abhandl.  d.  deutsch.  naturw.-med.  Vereines  f. 
Boehmen  "Lotos,"  11,  heft  4,  52-58. 

370.  ■   1909.  Neue  Andriasreste  aus  dem  Tonen 

von  Preschen  bei  Bilin.    "Lotos,"  lvii,  6. 

371.  Lea,    Isaac.  1849.  On   traces   of   a   fossil   reptile 

(Sauropus  primaevus)  found  in  the  old  Red 
Sandstone  (Pennsylvania).  Rep.  Brit.  Assn.  Adv. 
Sci.,  56,  2d  part;  Proc.  Amer.  Phil.  Soc,  91-94. 

372.  .  1855.  Fossil  footmarks  in  the  Red  Sand- 
stone of  Pottsville.  1-16,  with  a  large  plate. 
Large  folio;  Trans.  Amer.  Phil.  Soc,  x,  1852,  Apr. 

1-13.  P's.  31-33- 

373.  Leidy,  Joseph.   1856.  Notices  of  remains  of  extinct 

vertebrated  animals  discovered  by  Professor  E. 
Emmons.  Proc.  Acad.  Nat.  Sci.  Phila.,  256. 
(Dictyocephalus  elegans.) 

374.  .  1878.  Fossil  foot-tracks  of  the  Anthracite 

Coal  Measures.  Proc.  Acad.  Nat.  Sci.  Phila., 
xxxi,  164-165. 

375.  Le   Roy,   J.   J.  1874.  Notiz  ueber  Archegosaurus 

decheni  Goldfuss  und  A.  latirostris  Jordan. 
Niederl.  Archiv.  f.  Zoologie.  n,  89-98. 

376.  Lesley,  J.  P.  1889.  A  dictionary  of  the  fossils  of 

Pennsylvania  and  neighboring  states.  In  three 
volumes.    Published  by  the  Penn.  Geol.  Sun'. 

377.  Leunis,  Johannes.  1883.  Synopsis  der  Thierkunde. 

3d  Auflage,  Bd.  I,  599-631.    (Amphibia.) 

378.  Lillie,  Frank  R.  1908.  The  development  of  the 

chick.  The  Vertebrce,  412-424,  figs.  246,  249, 
250. 

379.  Lloyd,  G.  1849.  On  a  new  species  of  Labyrintho- 

don  from  the  New  Red  Sandstone  of  Warwick- 
shire. Rept.  Brit.  Assn.  Adv.  Science,  56.  (See 
Huxley  (328)  and  F.  von  Huene  (324).) 

380.  Logan,  William.  1842.  Proc.  Geol.  Soc,  London 

hi,  pt.  11,  707. 

381.  Lohest,  Maximin.   1888.  Decouverte  de  plus  an- 

cien  Amphibien  connu  et  de  quelques  Fossiles 
remarquables  de  Famennien  supeneur  de  Modave. 
Notice  preliminaire.  Ann.  de  la  Soc  Geol. 
Belgique,  XV,  p.  cxx,  fig.  r. 


382. 
383. 

384. 
385. 

386. 

387. 
388. 

389. 

390 

391- 
392- 

393- 

394- 

395- 

396. 

397- 

398- 
399- 

400. 


Lortet,  Dr.  L  1892.  Les  Reptiles  fossiles  du  Bassin 
du  Rhone.  Ordre  des  Amphibiens,  125-127,  pi. 
xii;  Archives  du  Museum  d'Hist.  Natur.  de 
Lyon,  v. 

Lucas,  F.  A.  1904.  A  new  Batrachian  and  a  new 
reptile  from  the  Trias  of  Arizona.  Proc.  U.  S- 
Nat.  Mus.,  xxvii,  193-195,  pi.  (Heterodonto- 
suchus,  a  belodont;  Metoposaurus  fraasii,  a 
labyrinthodont.) 

I. ydkkkkr,  Richard.  1879.  Reptilia  and  Batrachia 
(Amphibia).  Paleontologia  Fndica,  ser.  iv,  1, 
1-36,  pis.  i-vi;  Amphibia,  17-20,  pis.  iii-vi. 
(Pachygonia,  Gonioglyptus,  Mastodonsaurus.) 

.  1880.  A  sketch  of  the  history  of  the  fossil 

Vertebrataof  India.  Jour.  Asiatic  Society  Bengal, 
xlix,  pt.  11,  8-40;  Fossil  Batrachians,  16,  list  of 
species,  38-39. 

1 88 1.  Note    on    some    Gondwana    verte- 


brates.  Rec  Geol.  Surv.  India,  xiv,  174.    (Gonio- 
glyptus, Pachygonia,  Glyptognathus.) 

-.  1883.  Note  on  the  Bijori  Labyrinthodont. 


Rec.  Geol.  Surv.  India,  xvi,  93. 

1883.  Synopsis  of  the  fossil  Vertebrata  of 


India.    Rec.  Geol.  Surv.  India,  xvi,  61 ;  Amphibia, 
64,  88. 

1885.  The  Reptilia  and  Amphibia  of  the 


Maleri  and  Denwa  Groups.    Paleontologia  Indica, 
ser.  iv,  1,  pt.  5,  6  pis. 

-.  1885.  The  Bijori  Labyrinthodont.    Paleon- 


tologia Indica,  ser.  iv,  I,  1-14,  pt.  4;  also,  Cope, 
Amer.  Nat.,  xix,  592. 

-.  1887.  The  fossil  Vertebrata  of  India.    Rec. 


Geol.  Surv.  India,  xx,  p.  68. 

-.  1887.  Dr.   Hermann  Credner  on  the  de- 


velopment of  Branchiosaurus.    Geol.  Mag.,  n.  s., 
Dec  ill,  iv,  276-278. 

-.  1890.  Catalogue  of  fossil  Amphibia  and 


Reptilia  in  the  British  Museum  (Natural  History). 
Amphibia,  part  iv,  121-225. 

1891.  On  a  Labyrinthodont  skull  from  the 


Kilkenny  Coal  Measures  (Ireland).  Quart.  Jour. 
Geol.  Soc,  xlvii,  343-347.  1  fig- 

.  The  new  natural   history,     v,  Amphibia, 

257-310;  Labyrinthodontia,  311.  (See  also 
Nicholson  and  Lydekker  (499).) 

Lyell,  Chas.,  and  J.  W.  Dawson.  1853.  On  the 
remains  of  the  reptile  (Dendrerpeton  acadianum) 
and  a  land  shell  (Coal  Measures).  Quart.  Jour. 
Geol.  Soc,  ix,  58-63,  with  pis. 

Maggi,  Leopoldo.  1898.  Placche  osteodermiche 
interparietali  Stegocephali  e  rispondenti  Centri 
Ossificazione  interparietali  dell  'Uomo.  Reale 
Inst.  Lombard,  di  Sci.  Lett.  (2),  xxxi,  3X1,  228, 
pi.  1. 

.  1897.  Resultats   de    Recherches    morpho- 

logiques  sur  des  Os  et  des  Fontanelles  du  Crane 
humain.    Arch.  Ital.  d.  Biol.,  T.  27,  230-238. 

Makowsky,  Alex.  1876.  Ueber  einen  neuen  Laby- 
rimhodonten —  Archegosaurus  austricus.  Sitz- 
ungsb.  d.  lb  Akad.  wiss.  Wien.  Math.  Xatur. 
Classe,  Abth.  1.  lxxiii,  Heft  3,  155-166. 

und   A.   Rzehak.    1884.    Die  geologischen 

Verhaltnisse  der  Umgebung  von  Brunen.  Ver- 
handlgn.  d.  Nat.  Verein  zu  Brunn,  xxn.  (Die 
fruhere  Archegosaurus  austricus  wird  zu  Melaner- 
peton  gestellt.) 


212 


THE   COAL   MEASURES  AMPHIBIA   OF   NORTH   AMERICA. 


401.  Malbranc,  M.  1875.  Von  der  Seitenlinie  und  ihre 

Sinnesorganen  bei  Amphibien.  Leipzig.  Zeit. 
f.  wiss.  Zool.,  xxvi,  24,  pis.  1-4. 

402.  Mantell,    G.  A.     1 85 1.     Petrifactions    and   their 

teachings.    Fossil  Batrachians,  183. 

403.  .     1866.    Wonders  of  geology.    Labyrintho- 

donts,  551. 

404.  Marsh,  O.    C.   1862.  Discovery   of   new   Enalio- 

saurian  remains  in  the  Coal  Formation  of  Nova 
Scotia.  On  the  saurian  vertebra?  from  Nova 
Scotia.  Amer.  Jour.  Sci.  and  Arts,  xxxiv,  Mar.  I, 
pi.  1,  figs.  1-2.  See  also  Neues  Jahrb.  Mineral., 
Geol.  u.  Paleon.,  1863,  247;  Canad.  Nat.  and 
Geol.,  vii,  205-213;  Quart.  Jour.  Geol.  Soc. 
London,  xix,  52-56;  Huxley,  Scientific  Memoirs, 
n,  566. 

405.  .  1887.  Eobatrachus    agilis   in    the    (Como 

Beds)  Jura  of  North  America.  Amer.  Jour.  Sci., 
xxxm,  328;  Vertebrate  fossils  of  the  Denver 
Basin,  Monograph  U.  S.  Geol.  Surv.,  xxvii,  508, 
1897. 

406.  .  1894.  Footprints  of  vertebrates  in  the  Coal 

Measures  of  Kansas.  Amer.  Jour.  Sci.,  xlviii, 
2d  ser.,  July-Dec,  81-84,  pis.  ii-iii. 

407.  .  1896.  Amphibian  footprints  from  the  De- 
vonian. Amer.  Jour.  Sci.  (4),  11,  374-375,  with 
fig- 

408.  Matthew,  G.  F.  1903.  New  genera  of  Batrachian 

footprints  of  the  Carboniferous  system  of  eastern 
Canada.  Canadian  Record  of  Science,  ix,  No.  2, 
99-1 11,  with  figs. 

409-  •  I903-  An   attempt    to    classify    Paleozoic 

Batrachian  footprints.  Trans.  Roy.  Soc.  Canada 
(2),  ix  (m),  1903,  109-121. 

4io-  •  I9°3-  On  Batrachian  and  other  footprints 

from  the  Coal  Measures  of  Joggins,  N.  S.  Bull. 
Nat.  Hist.  New  Brunswick,  v,  No.  xxi,  103-108, 
with  plate  of  excellent  photos;  No.  xxn,  247. 

4ii-  •  I905-  Note    in    reference    to    Batrachian 

footprints.    Bull.  Soc.  New  Brunswick,  xxi,  102. 

412-  •  '9°5-  Note    on    the    genus    Hylopus    of 

Dawson.  Bull.  Soc.  New  Brunswick,  xxn,  247- 
252,  figs. 

4'3-  •  1904-  New  species  and  a  new  genus  of 

Batrachian  footprints  of  the  Carboniferous  sys- 
tem in  Eastern  Canada.  Trans.  Roy.  Soc. 
Canada,  x  (iv),  77-110,  pis.  1-6. 

414.  Matthew,  W.  D.  1909.  The  oldest  land  reptiles  of 
North  America.  Amer.  Mus.  Jour.,  ix,  No.  4, 
91-95.  photos  of  Eryops  and  Diplocaulus. 

4'5-  •  1911-  The  Amphibians  of  the  great  Coal 

Swamp.  Amer.  Mus.  Jour.,  xi,  No.  6,  197-200. 
(Restoration  of  Eryops.) 

416.  Merrill,  Geo.  P.  1907.  Catalogue  of  the  type 
and  figured  specimens  of  fossils,  minerals,  rocks 
and  ores.  Part  II:  Fossil  Vertebrates,  9-12' 
Bull.  U.  S.  Nat.  Mus.,  No.  53,  pt.  11.  Two  species 
of  Amphibia  are  listed  as  fishes,  74,  77 

417-  Metcalfe,  A.  T.  1884.  On  further  discoveries  of 
vertebrate  remains  in  the  Triassic  strata  of  the 
South  Coast  of  Devonshire,  between  Budleigh 
Salterton  and  Lidmouth.  Quart.  Jour.  Geol' 
Soc,  XL,  257. 

418.  Meyer,  Hermann  von.  1838.  Recherches  sur  les 
Ossem.  foss.  du  gres  bigarre  de  Soultz  les  Bains. 

4 ''■  •  l834-  Odontosaurus  voltzii.     Mem.  de  la 

Soc.  d'hist.  nat.  de  Strasbourg. 


420.  Meyer,  Hermann  von.   1842.   Labyrinthodontia. 

Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  301- 
304.   (Proposes  system  of  classification.) 

421.  .  1844.  Beitrage  zur  Paleontologie  Wurtem- 

bergs,  1-41,  mit  12  tafeln.  Enthaltend  die  fos- 
silen  Wirbelthierereste  aus  den  Triasgebilden  mit 
besonderer  Rucksicht  auf  die  Labyrinthodonten 
des  Keupers. 

422.  .  1844.  Apatheon  pedestris.     Neues  Jahrb. 

f.  Mineral.,  Geol.  u.  Paleon.,  336.  (He  regarded 
Apatheon  as  a  salamander-like  form,  a  Branchio- 
saurian.) 

423-  •  1845.  System  der  fossilen  Saurier.    Neues 

Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  278-285. 

424- •  1847.  Die  Steinbrucke  von  Oeningen.  Fossile 

Batrachier.  Fauna  der  Vorwelt,  Bd.  1,  18-40, 
pis.  9-10. 

425-  •  1847-1855.  Die  Saurier  des  Muschelkalkes 

mit  Rucksicht  auf  die  Saurier  aus  Bunten  Sand- 
stein  und  Keuper.  Fauna  der  Vorwelt,  Bd.  11, 
1-164,  pis.  58-64.  Large  folio.  (Mastodon- 
saurus,  Metopias,  Capitosaurus.) 

426. .  1848.  Apateon    pedestris    aus   der    Stein- 

kohlenformation  von  Mitnsterappel.  Paleonto- 
graphica, Bd.  1,  153-154,  pi.  xx,  fig.  1. 

427-  •  1848.  Mittheilung    an    Professor    Bronn. 

Labyrinthodontia.  Neues.  Jahrb.  f.  Mineral., 
Geol.  u.  Paleon.,  469.     (Labyrinthodon  ocella.) 

428.  — .  1849.  Ueber  den  Archegosaurus  der  Stein- 

kohlenformation.  Paleontographica,  Bd.  1,  209, 
Dec.  (1851),  pi.  33. 

429-  .  1851.  Mittheilung    an    Professor    Bronn, 

Rana  jaegeri.  Neues  Jahrb.  f.  Mineral.,  Geol.  u. 
Paleon.,  78. 

43°-  ' ■  1 85 1.  Beschreibung    der    fossilen     Deca- 

poden,  Fische,  Batrachier,  und  Saugethiere  aus 
dem  Tertiaren  Susswassergebilden  des  nordlichen 
Boehmens.    Paleontographica,  Bd.  11,  43. 

43  •■  •  1852.  Mittheilungen  an  Professor  Bronn. 

Neues  Jahrb.  f.  Mineral,  Geol.  u.  Paleon.:  Pateo- 
batrachus  gigas,  465;  Rana  noggerathi,  58;  R. 
troscheli,  466;  R.  salzhauserensis,  467. 

432.  .  1853.  Mittheilungen  an  Professor  Bronn. 

Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.:  Rana 
meriani,  163,  185;  Pakeobatrachus  gigas,  162. 

433-  •    1854.    Mittheilung  ueber  Archegosaurus. 

Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  424. 

434-  •  1856.  Ueber  Osteophone  roemeri,  Briefl. 

Mittheil.  an  Professor  Bronn.  Neues  Jahrb.  f. 
Mineral.,  Geol.  u.  Paleon.,  824. 

435-  •  1857-  Mittheilung    an    Professor    Bronn. 

Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.:  Pakeo- 
batrachus gracilis,  555. 

436.  ■ — .  1857.  Reptilien  aus  der  Steinkohlenforma- 

tion  in  Deutschland.  Paleontographica,  Bd.  vi, 
219-220,  text-fig.    (Archegosaurus.) 

437- ■  1857.  Osteophorus  roemeri,  Beschreibung 

derselben  in  Roemer:  Ueber  Fisch-  und  Pflanzen- 
fuhrende  Mergelschiefer  des  Rothliegenden  bei 
Neundorf.    Zeit.  d.  deutsch.  geol.  Gesell.,  ix,  61. 

438.  .  1858.  Reptilien  aus  der  Kohlsteinforma- 

tion  in  Deutschland.  Cassel,  Folio;  also  Pale- 
ontographica, Bd.  VI,  59-218,  Tafeln  viii-xxiii. 
(Archegosaurus.) 

439-  •  1858.  Labyrinthodonten  aus  dem  bunten 

Sandstein  von  Bernburg.  Paleontographica,  Bd. 
vi.  221-245,  Tafeln  xxiv-xxvii. 


BIBLIOGRAPHY   OF    THE    FOSSIL   AMPHIBIA. 


213 


440.  Mkyer,  Hermann  von.  1858.  Mittheilung  an 
Professor  Bronn.  Neues  Jahrb.  f.  Mineral.  Geol. 
u.  Paleon.:  Rana  danubina,  203. 

44'-  ' — ■  1859-  Mittheilung    an    Professor    Bronn. 

Neues  Jahrb.  f.  Mineral.  Geol.  u.  Paleon.:  Triton 
basal ticus,  430;  Andrias  tschudii,  723. 

442.  ■  i860.  Ueber  Archegosaurus.     Erwiderung 

auf  die  Ausfuhrungen  von  Quenstedt  in  "  Epochen 
der  Natur,"  p.  410.  Brierl.  Mittheil.  an  Professor 
Bronn.  Neues  Jahrb.  f.  Mineral.  Geol.  u.  Paleon., 
71-72. 

i860.  Osteophorus  roemeri  aus  dem  Roth- 


443 


444 


445 


446 


447 


448 


449 


450 


45i 


452 


453 


454- 


liegenden    von    klein    Neundorf    in    Schliesen. 

Paleontographica,  Bd.  Vll,  99-104,  pi.  xi. 
.  i860.  Frocsche     aus     Tertiar     Gebilden 

Deutschlands.     Paleontographica,  Bd.  vn,  123- 

182,  pis.  16-22. 
.  i860.  Melosaurus  uralensis  aus  der  Per- 

mische  System  des  westlichen  Urals.    Paleonto- 
graphica, Bd.  vh,  90-98,  pi.  x. 
.  i860.  Salamandrinen  aus  der  Braunkohle 

am  Rhein  und  in  Boehmen.     Paleontographica, 

Bd.  vn,  47-73,  pis.  viii,  ix. 
.  1863.  Heliarchon    furcillatus,    ein    Batra- 

chier  aus  der  Braunkohle  von  Rott  in  Siebenge- 

birge.     Paleontographica,  Bd.  x,  292-298,  Taf. 

50,  figs.  5-6. 

1866.  Reptilien  aus  dem  Kupfersandstein 


Russlands.    Paleontographica,  Bd.  xv,  97-130. 
Miai.l,  L.  C.  1873.  Report  on  the  Labyrinthodonts 

of  the  Coal  Measures.     Rept.  Brit.  Assn.  Adv. 

Sci.,  225,  pis.  1-3. 
.  1874.  On   the  classification  of  the  Lahy- 

rinthodontia.    Rept.  Brit.  Assn.  Adv.  Sci.,  149- 

192,  with  2  plates. 

1874.  The  classification  of  the  Labyrintho- 


donts.   Geol.  Mag.,  2d  ser.,  1,  513. 

1874.  On  the  occurrence  of  a  Labyrintho- 


dont   in   the  Yoredale   Rocks   of   Wensleydale. 
Quart.  Jour.  Geol.  Soc,  xxx,  775. 

-.  1874.  On  the  remains  of  Labyrinthodontia 


from  the  Keupcr  Sandstone  of  Warwick,  pre- 
served in  the  Warwick  Museum.  Quart.  Jour. 
Geol.  Soc,  xxx,  417-435,  pis.  26-28.  (Masto- 
donsaurus  pachygnathus;  Labyrinthodon  lepto- 
gnathus;  Diadetognathus  varvicensis.) 

1875.  Sur  les  Labyrinthodontes  du   Ter- 


rain houiller.    Deuxieme  partie.    Jour,  de  Zool., 
iv,  I9-37- 

455.  Miller,  S.  A.  1889.  North  American  geology  and 

paleontology.    Amphibia,  614. 

456.  Mivart,  Geo.  1870.  On  the  axial  skeleton  of  the 

Urodela.    Proc.  Zool.  Soc.  London,  260-278. 

457.  Moodie,  Roy  L.  1908.  Dissorophus — A  correction. 

Science,  n.s.,  xxvn,  No.  679,  30. 

-.  1908.  The  lateral  line  system  in  extinct 


458. 


459- 


460. 


461. 


Amphibia.   Jour.  Morphol.,  xix,  No.  2,  511-540, 
17  figs. 
.  1908.  The  ancestry  of  the  caudate  Am- 
phibia.   Amer.  Nat.,  42,  361-373,  figs.  1-10. 
-.  1908.  The  dawn  of  quadrupeds  in  North 


America.    Pop.  Sci.  Monthly,  lxxii,  June,  558- 
566,  5  figs. 

1908.  The  clasping  organs  of  extinct  and 


recent  Amphibia.    Biol.  Bull.,  xiv,  No.  4,  Mar., 
249-259,  with  5  figs. 


462. 
463. 

464. 
465- 

466. 
467. 

468. 
469. 

470. 
47i- 

472. 
473- 

474- 

475- 

476. 

477- 

478. 

479- 

480. 
481. 
482. 

483- 
484. 


Moodie,  Roy  L.  1909.  New  forms  (extinct  Amphi- 
bia) from  the  Carboniferous.  Jour.  Geol.,  xvn, 
No.    1,  Jan.-Peb.,  38-82,  24  figs. 

.  1909.  The    Carboniferous    quadrupeds  of 

Kansas,  Ohio,  Illinois,  and  Pennsylvania  in  their 
relation  to  the  classification  of  the  so-called 
Amphibia  and  Stegocephalia.  Trans.  Kans. 
Acad.  Sci.,  xxii,  239-243,  pis.  1-3. 

.  1909.  Carboniferous  air-breathing  verte- 
brates of  the  United  States  National  Museum. 
Proc.  U.  S.  Nat.  Mus.,  37,  ri-28,  pis.  4-10. 

1909.  New  or  little-known  forms  of  Car- 


boniferous Amphibia  in  the  American  Museum 

of   Natural   History.     Bull.   Amer.   Mus.    Nat. 

Hist.,  xxvi,  art.  xxv,  347-357,  pis.  lviii-lxv. 
.  1909.  The    Lysorophidie.      Amer.     Nat., 

XLiii,  116.     (Review.) 
.    1909.    The    Stegocephala.    Amer.     Nat., 

XLiii,    119.     (A    review    of    paper    by    Hugo 

Schwarz.) 

-.  1909.  The  morphology  of  the  vertebrate 


sacral  rib.    Anat.  Anz.,  xxxiv,  No.  15,  361-364. 
-.  1909.  The  Microsauria,  ancestors  of  the 


Reptilia.  Geol.  Mag.,  Dec.  v,  vi,  No.  539, 216-220, 
1  fig. 

1910.  The  Amphibia  of  the  Mazon  Creek 


Shales.    Science,  n.  s.,  xxxi,  No.  789,  p.  233. 

1910.  The  alimentary  canal  of  a  Carbonif- 


erous salamander.    Amer.  Nat.,  XLIV,  No.  522, 

367-375.  4  figs- 

-.  1910.  A  new  Labyrinthodont  from  Kansas. 


Science,  n.  s..  xxxii,  No.  829,  Nov.  18. 

-.  191 1.  A    new    Labyrinthodont    from    the 


Kansas  Coal  Measures.    Proc.  U.  S.  Nat.  Mus., 

xxxix,  489-495,  4  figs. 
.  191 1.  Two  Amphibians,  one  of  them  new, 

from  the  Carboniferous  of  Illinois.     Proc.  U.  S. 

Nat.  Mus.,  XL,  429-433,  figs.  1-2. 
-.  191 1.  Recent  contributions  to  a  knowledge 

of  the  extinct  Amphibia.    Amer.  Nat.,  xi.v,  June, 

375-384- 

-.  191 1.  The     temnospondylous     Amphibia, 


and  a  new  species  of  Eryops  from  the  Permian  of 
Oklahoma.  Kans.  Univ.  Sci.  Bull.,  v,  No.  13, 
235-253.  pis.  xlix-liv. 

-.  1912.  The  skull  structure  of  Diplocaulus 


magnicornis  and  the  amphibian  order,  Diplo- 
caulia.  Jour.  Morphol.,  xxm,  No.  1,  31-39,  pis. 
1-2. 

-.  1912.  The  Pennsylvanic  Amphibia  of  the 


Mazon  Creek,  Illinois,  Shales.    Kans.  Univ.  Sci. 
Bull.,  vi,  No.  2,  323-359.  Pis.  1-14- 

-.  1912.  The  Mazon  Creek,  Illinois,  Shales 


and  their  amphibian  fauna.     Amer.  Jour.  Sci., 
xxxiv,  Sept.,  277-285,  figs.  1-2. 

1912.  An  American  Jurassic  frog.    Amer. 


Jour.  Sci.,  xxxiv,  Sept.,  286-288. 

1914.  The  fossil  frogs  of  North  America. 


Amer.  Jour.  Sci.,  xxxvm,  Dec,  53i-53<>.  figs-  1-2. 
-.  1914.  A  list  of  the  described  species  of 


fossil  Amphibia.    Kans.  Univ.  Sci.  Bull.,  9,  No.  2, 
13-28. 

1915.  A  remarkable   Microsaur  from  the 


Coal  Measures  of  Ohio.    Science,  n.  s.,  xli.  No. 
1044,  34-35- 

.  1915.  Coal  Measures  Amphibian  with  an 

osseous  tarsus.     Amer.  Jour.  Sci.,  xxxix.  May, 
509-512,  figs.  1,  2. 


214 


THE    COAL   MEASURES    AMPHIBIA   OF    NORTH    AMERICA. 


485.  Moodie.RoyL.  igij.ThescaledAmphibiaoftheCoal 

Measures.  Science,  n.  s.,  xli,  No.  1056,  463-464. 

486.  — .   19 1 5.  Some     recent     studies     on      fossil 

Amphibia.    Amer.  Nat.,  xux,  June,  369-376. 

487-  •  1915-  A   new   fish   brain   from   the   Coal 

Measures  of  Kansas,  with  a  review  of  other  fossil 
brains.    Jour.  Comp.  Neurol.,  xxv,  No.  2,  164. 

488. .  1915.  A  further  contribution  to  a  knowledge 

of  the  lateral  line  system  in  extinct  Amphibia.  Jour. 
Comp.  Neurol.,  xxv,  No.  4,  317-328,  figs.  1-9. 

.  1915.  Some   methods   of   studying   fossils 

embedded  in  coal.    Kans.  Univ.  Sci.  Bull.,  IX, 
No.  16,  187-193,  figs.  1-3,  pi.  l. 

.  The  migrations  and  geographic  distribu- 
tion of  the  fossil  Amphibia.  Amer.  Jour.  Sci., 
xl,  186-190,  1  map. 

■  The  Coal   Measures  Amphibia   and  the 

Crossopterygia.    Amer.  Nat.,  xlix,  637-644. 


489. 


489a. 


4896. 


490.  Mudge,  B.F.  1873.  Recent  discoveries  of  fossil  foot- 

prints in  Kansas.    Trans.  Kans.  Acad.  Sci. ,  11,  7-9. 

491.  Munster,    Georg,    Graf    zu.    1834.     Vorlaufige 

Nachricht  ueber  einige  neuen  Reptilien  im 
Muschelkalk  von  Baiern.  Neues  Jahrb.  f.  Min- 
eral., Geol.  u.  Paleon.,  527.     (Mastodonsaurus.) 

492-  •    1836.    Capitosaurus     arenaceus.       Neues 

Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  580. 

493-  •  1839-  Mastodonsaurus  adriani.     Beitrage 

zur  Petrefactionkunde,  102. 

494.  Murchison,  Roderick  I.  1872.  Siluria— A  history 

of  the  oldest  rocks  in  the  British  Isles  and  other 
countries.    5th  ed.,  302,  343. 

495.  Newberry,  J.  S.  1856.  Description  of  several  new 

genera  and  species  of  fossil  fishes  from  the  Car- 
boniferous strata  of  Ohio.  Proc.  Acad.  Nat.  Sci. 
Phila.,  viii,  98. 

496-  — ■ •  1867.  On  some  fossil  reptiles  and   fishes 

from  the  coal  strata  of  Ohio,  Kentucky,  and 
Illinois.     Proc.  Amer.  Assn.  Adv.  Sci.;  144-146. 

497-        •  1878.  Geology  of  Jefferson  County,  Ohio. 


Geol.  Surv.  Ohio,  in,  736-737. 

498- •   1889.  Palaeozoic  fishes  of  North  America. 

Monograph  U.  S.  Geol.  Surv.,  xvi,  228.  (Dis- 
cusses Kammplatten;  Amphibia  of  Linton,  Ohio, 
211-214;  Fauna  of  Mazon  Creek,  214.  See  also 
Hodge,  1878,  Geol.  Surv.  Ohio,  in,  592.) 

499.  Nicholson  and  Lydekker.  1889.  Manual  of  pale- 

ontology.   Amphibia,  II,  1018-1045. 

500.  Oldham,  Dr.  R.  D.  1875.   On  a  salamandriform 

Labyrinthodont  from  the  Coal  Measures  near 
Castlecomer,  Ireland.  Rept.  Rugby  School  Nat. 
Hist.  Soc,  74-76,  pi.  vii. 

501.  — - — .  1893.  Geology   of    India.      2d    ed.      The 

Gondwana  System,  149. 

502.  Owen,   Sir   Richard.      1 840-1 845.     Batrachians. 

Odontography,  187-218,  pis.  62,  63,  64. 

503-  •  1842.  On  the  teeth  of  the  species  of  the 

genus  Labyrinthodon  of  Jaeger  from  the  German 
Keuper  and  the  sandstone  of  Warwick  and  Leam- 
ington. Ann.  and  Mag.  Nat.  Hist.,  vm,  58;  also, 
Geol.  Trans.  11,  ser.  vi,  503. 

504-  •  1842.  Description  of  parts  of  the  skeleton 

and  teeth  of  the  genus  Labyrinthodon.  with  re- 
marks on  Cheirotherium.  Trans.  Geol.  Soc. 
London,  vi,  pt.  2,  515-543,  pis.  xliii-xlvii. 

5»5-  •   1842.  On  the  teeth  of  species  of  the  genus 

Labyrinthodon.  Trans.  Geol.  Soc.  London,  vi, 
pt.  2,  503-513,  pi.  43,  fig.  j,  (Discusses  validity 
of  term  Mastodonsaurus.    Read  Jan.  11,  1841.) 


506 

507 
508. 

5°9- 
510. 

5"- 
512. 


513- 
514- 


.  Owen,  Sir  Richard.  1847.  On  the  Batracholites  in- 
dicative of  a  small  species  of  frog  (Rana  pusilla). 
Quart.  Jour.  Geol.  Soc.,  m,  224.  (See  Clark,  103, 
p.  221.) 

.   1853.  Notice  of  a  batrachoid  fossil  in  the 

British    Coal    Shale.     Quart.    Jour.   Geol.   Soc. 
London,  ix,  67-70,  pi.  iii. 

-.  1853.  Notice  of  a  batrachoid  fossil —  Para- 


515- 

516. 
517- 

518- 

519- 
520. 


batrachus  colei — in  the  British  Coal  Shale.  Quart. 
Jour.  Geol.  Soc.  London,  x,  207. 

•  1853.  Baphetes  planiceps — a  fossil  em- 
bedded in  a  mass  of  coal  from  Nova  Scotia. 
Quart.  Jour.  Geol.  Soc.  London,  x,  207-208,  pi.  ix. 

.  1854.  Description    of    the    cranium    of   a 

Labyrinthodont  reptile  (Brachyops)  from  the 
Triassic  of  Central  India.  Quart.  Jour.  Geol. 
Soc.  London,  x,  473.     (Abstract.) 

•  1855.  Description    of    the    cranium    of   a 

Labyrinthodont  reptile  from  Mangala,  Central 
India.     Quart.  Jour.  Geol.  Soc.  London,  xi,  37,  pi. 

.  1859.  On  the  orders  of  fossil  and  recent 

Reptilia  and  their  distribution  in  time.  Rept. 
Brit.  Assn.  Adv.  Sci.,  153-166. 

.   1861.  Paleontology,  2d  ed.,  Amphibia,  193. 

.   1862.  Descriptions  of  specimens  of  fossil 

Reptilia  discovered  in  the  Coal  Measures  of  the 
South  Joggins,  Nova  Scotia.  Quart.  Jour.  Geol. 
Soc.  London,  xvm,  238-244,  pis.  9-10. 

-.  1865.  Description  of  some  remains  of  an 


522, 
523. 
524- 

525- 
526. 

527- 


air-breathing   vertebrate   (Anthrakerpeton   cras- 

sosteum)  from  the  Coal  Shale  of  Glamorganshire. 

Geol.  Mag.,  11,  6-8,  pis.  i-ii. 
.  1866.  On  the  anatomy  of  the  Vertebrata. 

Amphibia,  I,  8. 
■  1876.  On   Petrophryne   granulata    Ow.,  a 

Labyrinthodont  reptile  from  the  Trias  of  South 

Africa.     Bull,  de  la  Soc.  Imp.  de  Nat.  de  Moscou. 

(Review.) 
.   1876.  Catalogue  of  the  fossil   Reptilia  of 

South    Africa   in   the   collection   of    the    British 

Museum.     Labyrinthodontia,  67-70,  pi.  xx,  figs. 

13-20;  pi.  lxx,  fig.  3. 
.  1884.  On    a    Labyrinthodont    Amphibian 

from  the  Cape  of  Good  Hope.    Quart.  Jour.  Geol. 

Soc.  London,  XL,  333,  2  pis. 
.   1884.  On    Rhytidosteus    capensis    Ow.,  a 

Labyrinthodont  from  the  Trias  of  Cape  of  Good 

Hope.    Ann.  and  Mag.  Nat.  Hist.  (5),  xin,  481. 
Pabst,  Wm.   1896-1905.  Beitrage  zur  Kenntniss  der 

Thierfahrten  in  dem  Rothliegenden  Deutschlands. 

Zeit.  der  deutsch.  geol.  Gesell.,  1896,  xlviii,  638, 

808;  1900,  lii,  48;  lvii,  1 -14,  361-379,  with  pis. 

(Some  of  the  tracks  described  may  be  amphibian.) 
Pander,  Chr.  H.  1858.  Ueber  die  Ctenodipterinen 

des  devonischen  Systems. 
Pictet,  F.  J.  1853.  Traite  de  paleontologie,  2d  ed., 

tome  premier. 
Poli.ard.H.B.  1891.  On  the  anatomy  and  phylogen- 

etic  position  of  Polypterus.  Anat.  Anz. ,  VL338 ;  Zool. 

Jahrb.(Morphol.Abth.),v,  1893, 387-428',  with  pis. 
Portis,  A.   1885.  Resti  di   Batraci  fossili   Italiani: 

appunti  paleontologi,  pt.  2,  p.  30. 
Pusch,    Bergrath.   1842.  Fossile   Batrachier   und 

Ophidier  Rcste  aus  Podolien.     Neues  Jahrb.  f. 

Mineral.,  Geol.  u.  Paleon.,  179-180. 
Quenstedt,  F.  A.  von.   1850.  Die  Mastodonsaurier 
im  gruenen  Keupersandstein  Wiirtembergs  sind 
Batrachier,  4to,  1-34,  4  Tafeln. 


BIBLIOGRAPHY    OF   THE   FOSSIL   AMPHIBIA. 


31! 


528.  Qualen,  Wangenheim  von.  1S45  and  1852.  Two 

papers  on  the  skull  of  Zygosaurus  lucius.  Bull. 
Soc.  Imp.  Nat.  Moscou,  xvm,  389  (1852), 
47-!- 

529.  .  1 861.  Bemcrkungen    zum    Archegosaurus. 

Neues  Jahrb.  f.  Mineral.  Geol.  u.  Paleon.,  294-300, 
with  pi.  iii.  (See  also  Baur,  1886,  Amer.  Nat.,  173, 
and  Zool.  Anz.,  1886,  Nr.  216.) 

53«-  •  1885.  Atlas  zum  Handbuch  der  Petrifac- 

tionskunde.     Amphibia,  pi.  18. 

531.  Raymond,  Percy  E.   1907.  On  the  discovery    of 

reptilian  remains  in  the  Pennsylvanian  near 
Pittsburgh,  Pa.  Science,  n.  s.,  xxvi,  676,  835. 

532.  Rjabinin,   M.   A.  1911.  Debris  de  Stegocephales 

trouves  aux  Mines  de  Kargala  Gouvernement 
d'Orembourg.  Bull,  du  Comm.  Geol.  St.  Peters- 
burg, xxx,  No.  185,  pis.  1-2. 

533.  .  1912.  Ueber    die    Ueberreste    der    Stego- 

cephalen  aus  den  Kargalinokischen  Bergwerken 
des  Gouvernments  Orenberg,  St.  Petersburg, 
37  pages,  pis.  1-2. 

534.  Roemer,  Ferdinand.  1857.  Ueber  Fische  und  Pfian- 

zenfiihrende  Mergelschiefer  des  Rothliegenden 
bei  Klein-Neundorf  und  ueber  Acanthodes 
gracilis.     Zeit.  d.  deutsch.  geol.  Gesell.,  ix,  61. 

535.  Scheuchzer,  J.  J.  1726.  Part  of  two  human  skele- 

tons petrified.  Phil.  Trans.  Roy.  Soc.  I-ondon, 
1726,  abridged  edition,  vn  (1724-1734),  129. 
(Translated  from  the  Latin  of  a  letter  to  Sir  Hans 
Sloan,  a  member  of  the  Royal  Society.) 

536.  Schmid,     E.  1853.  Die     organischen     Reste     des 

Muschelkalks.  Neues  Jahrb.  f.  Mineral.,  Geol.  u. 
Paleon.,  15. 

537.  Schmidt,  M.  1907.  Labyrinthodontenreste  aus  dem 

Hauptkonglomerat  von  Altensteig  im  wurtem- 
bergischen  Schwarzwald.  Stuttgart.  Mittheil.  d. 
Geol.  Abth.,  d.  K.  Wurtemb.  Statest.  Landes- 
amts,  Jahrg.  63,  No.  2,  1-10,  with  plate. 

538.  Schonfeld,  G.  1911.  Branchiosaurus  tener  Schonf. 

Ein  neuer  Stegocephal  aus  dem  rothl.  des  Nord- 
west  Sachsens.    Isis,  19-43,  with  plate. 

539.  Schroder,    Henry.      1913.      Ein    Stegocephalen 

Schadel  von  Helgoland.  Jahresb.  d.  k.  preuss. 
geol.  Landesanstalt  fur  1912,  33,  Theil  II,  Heft  2, 
Taf.  15-21. 

540.  Schwarz,  Hugo.   1908.  Ueber  die  Morphologie  der 

Wirbelsaule  der  Tetrapoden.  Sitzungsb.  d. 
Gesell.  naturf.  Freunde  zu  Berlin,  315-329. 

541.  .  1908.  Ueber    die    Wirbelsaule    und    die 

Rippen  holospondyler  Stegocephalen  (Lepo- 
spondyli,  Zittel).  Beitrage  zur  Geol.  u.  Paleon. 
Oesterreichs-Ungarns  u.  des  Orients,  Bd.  xxi, 
Heft  1-2,  63-105,  36  figs,  and  Bibliography. 

542.  Scott,  W.  B.  1908.  An  introduction  to  geology.  2d 

revised  edition.  Amphibia,  608,  636,  652,  674, 
692. 

543.  Sedgwick,  Adam.  1905.  Stegocephali  in  "Student's 

Text-book  of  Zoology,"  11,  313. 

544.  Seeley,  H.  G.  1875.  On  the  posterior  portion  of  a 

lower  jaw  of  Labyrinthodon  (L.  lavisi)  from  the 
Trias  of  Lidmouth.  Quart.  Jour.  Geol.  Soc., 
xxxil,  278-284,  pi.  xix. 

545.  .  1879.  Amphibians  from  the  Permian  Rocks 

of  Bohemia.  Geol.  Mag.,  Dec.  11,  vi,  521-528,  4 
woodcuts;   1885,  80-87.     (A  review  of  Fritsch's 


work.) 


54&- 


1892.  Further  observation  on  Pareiasaurus. 


547- 
548. 
549- 

55<>- 
551. 

552- 
553- 

554- 

555- 

556. 

557- 

558. 
559- 

560. 
561. 

562. 

563 
564- 
565- 

566. 
567- 


Seeley,  H.G.  1907.  AnewLabyrinthodontfromthe 
Karoo  Beds,  Cape  Colony.  Geol.  Mag.,  Dec.  v, 
iv,  No.  x,  433-434  pi.  1. 

.  1908.  A   Labyrinthodonl  tooth  from   the 

upper  Karoo  Beds,  South  Africa.     Geol.  Mag., 
Dec.  v,  v,  No.  vi,  241-243,  pi.  x. 

Smith,  John.  Footprints  in  red  calciferous  sand- 
stones in  a  railway  cutting  2%  miles  N.  of 
West  Kilbride.  Trans.  Geol.  Soc.  of  Glasgow, 
ix,  201. 

Stejneger,  Lkonhakd.  1904.  Amphibia  versus 
Batrachia.    Science,  n.s.,  xx,  924. 

.  1907.  Herpetology  of  Japan  and  adjacent 

territory.    Bull.  No.  58,  U.  S.  Nat.  Mus.,  1-576, 
pis.  i-xxxv,  and  figs. 

Steinmann-Doderlein.  1890.  Elemente  der  Pale- 
ontologie.    Amphibia,  601. 

Steinmann,  Gustav.  1907.  Einfuhrung  in  die 
Paleontologie.  Zweite  Auflage.  2  Klasse:  Am- 
phibia, Lurche,  421-427,  figs. 

Stephens,  A.  T.  1886.  On  the  Biloela  Labyrinthc- 
dont.  Proc.  Linn.  Soc.  New  South  Wales,  u 
ser.,  1,  1 1 13. 

1888.  On  some  additional  Labyrinthodont 


fossils  from  the  Hawksbury  sandstone  of  New 
South  Wales.  Proc.  Linn.  Soc.  New  South  Wales, 
1  (2).  1175-1192. 

-.  1888.  Note   on   a    Labyrinthodont   fossil 


from  Cockatoo  Island  (Mastodonsaurus).    Proc. 
Linn.  Soc.  New  South  Wales  (2),  I,  931-940. 

1888.  Second   note  on    Platyceps  wilkin- 


soni.    Proc.  Linn.  Soc.  New  South  Wales,  11  (2), 

156-158,  pi.  1. 
Sternberg,  C.  H.  1901.  The  Permian  life  of  Texas. 

Trans.  Kans.  Acad.  Sci.,  94-98. 
Stevenson,  J.  J.  1907.  Carboniferous  of  the  Appa- 
lachian basin.    Bull.  Geol.  Soc.  Amer.,  xvm,  29- 

178. 
Storrie,  J.   1893-4.  Notes  on  a  tooth  of  a  species 

of  Mastodonsaurus.     Trans.  Cardif.  Nat.  Soc., 

xxvi,  pt.  11,  105,  pi. 
Stock,  Thomas.  1881.  On  some  British  specimens 

of  the  "Kammplatten"  or  "  Kammleisten "  of 

Professor  Fritsch.     Ann.  and  Mag.  Nat.  Hist., 

5th  ser.,  vm,  90-95,  pi.  vi. 
.  1882.  Notice  of  some  discoveries  recently 

made  in  Carboniferous  vertebrate  paleontology. 

Nature,  xxvn,   Nov.   2d,   22.     (Describes  new 

species  of  Ophiderpeton  (nanum)  and  Pteroplax 

(cornuta).) 
.     1882.     Further  observations  on   Kamm- 
platten and  Note  on  Ctenopthychius  pectinatus. 

Ann.   and    Mag.    Nat.   Hist.    (5),   ix,   253-257. 

pi.  viii. 
Strickler,  Ludwig.  1899.  Ueber  den   microscop- 

ischen  Bau  der  Faltenzahne  von   Eryops  mega- 

cephalus.     Paleontographica,  xlvi,  85-94,  1  fig., 

Taf  ein  xi-xii. 
Thevenin,     Armand.     1905.     Sur  lc    Decouvcrte 

d'Amphibiens  dans  le   Terrain  houiller  de  Com- 

mentry.   Comptes  Rendus  Acad.  Sci.,  cxli  (26), 

1268-1269. 
.  1906.  Amphibiens  et  Reptile  du  Terrain 

Houiller  de  France.     Annales  de  Paleontologie, 

Tome  1,  fas.  in,  1-19,  2  pis. 

1909.    Les   Stades  d'Evolution    des  plus 


Phil.  Trans.  Roy.  Soc.  London,  clxxxiii,  367. 


anciens  Quadruples  trouves  en  France.  Comptes 
rendus  des  Acad.  Sci.,  cxlix,  1222,  Dec.  20. 


216 


THE    COAL   MEASURES   AMPHIBIA   OF   NORTH   AMERICA. 


568.  Thevenin,  Armand.  1910.  Les  plus  anciens  Quad- 
ruples de  France.  Annales  de  Paleontogie,  Tome 
V,  1-64,  pis.  i-ix;  V,  fas.  1,  1-40,  pis.  1-6. 

569  Thompson,  James,  and  Professor  Young.  1869. 
On  new  forms  of  Pteroplax  and  other  Carbonifer- 
ous Labyrinthodonts  and  other  Megalichthys 
with  notes.  Rept.  Brit.  Assn.  Adv.  Sci.,  pt.  II, 
101. 

570.  Thyng,  F.  W.  1906.  The  squamosal  bone  in  Tet- 

rapodous  Vertebra ta.  Tufts  College  Studies,  n, 
No.  2,  scientific  series,  35-74. 

571.  Tomes,  Chas.  S.  1878.  A  manual  of  dental  Anat- 

tomy,  human  and  comparative.    Sth  ed.,  65. 

572.  Tomes,  John.  1850.  On  the  structure  of  the  den- 

tal tissues  of  the  order  Rodentia.  Phil.  Trans. 
Roy.  Soc.  London,  532.  (Remarks  on  the  struc- 
ture of  the  teeth  of  Labyrinthodon  jaegeri.) 

573.  Trautschold,  H.     1884.     Die    Reste  permischen 

Reptilien  des  Paleontologischen  Cabinets  der 
Universitat  Kasan.  Nouv.  Mem.  de  la  Soc.  Imp. 
Nat.  Moscou,  xv,  5-39,  Taf.  1-8. 

574.  Tschudi,  Johannes  J.  von.  1839.  Classification  der 

Batrachier  mit  Berucksichtigung  der  fossilen 
Thiere  dieser  Abtheilung  der  Reptilien.  Mem. 
de  la  Soc.  Scientif.  Nat.  de  Neuchatel,  4to,  11. 
Review:  Neues  Jahrb.  f.  Mineral.  Geol.  Paleon., 
1841,835-840. 

575.  .  1837.  Ueber  den  Homo  diluvii  Testis  A. 

Scheuchzer.  Neucs  Jahrb.  f.  Mineral.,  Geol.  u. 
Paleon.,  545~547- 

576.  Twelvetrees,  W.  H.  1880.  On  a  Labyrinthodont 

skull  (Platyops  ricardi  Twel.)  from  the  upper 
Permian  cupriferous  strata  of  Kargalinsk,  near 
Orenburg.  Bull,  de  la  Soc.  Imp.  des  Natl,  de 
Moscou,  lv,  No.  1,  117-122,  with  figs. 

577.  Udden,  J.  A.  1901.  Geology  of  Jefferson  Co.,  Iowa. 

Iowa  Geol.  Surv.,  xn,  Ann.  Rept.,  406. 

578.  Versluys,  J.     1909.     Die    Salamander    und    die 

ursprunglichsten  vierbeinigen  Land wirbel thiere. 
Naturwissenschaftliche  Wochenschrift,  Neue 
Folge,  Bd.  vih,  No.  3,  1-28,  figs.  1-5. 

579.  Vidal,  Louis  Mariano.  1902.  Sobre  la  presencia 

del  tramo  Kimeridgense  en  el  Montseih  y  hallazgo 
de  un  Batracio  en  sus  hiladas.  Memorias  Real 
Academ.  Cienc.  y  Artes  de  Barcelona,  iv,  No.  18. 

580.  .  1902.    Sur  le  presence  de  l'etage  Kime- 

ridgien  an  Montseih  (Province  de  Lerida,  Es- 
pagne)  et  decouverte  d'un  Batracien  dans  ces 
assises.  Memorias  Real  Academ.  Cienc.  y 
Artes  de  Barcelona,  iv,  No.  18. 

581 .  Vogt,  Carl.  1854.  Archegosaurus  und  all  Labyrintho- 

donten  sind  Amphibien  nicht  Reptilien.  Neues 
Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  676-677. 

582.  Wagner,  A.  1861.  Neue   Beitrage   zur   Kenntniss 

der  vorweltlichen  Fauna  des  lithograph. — Schie- 
fers.  Abhandl.  der  k.  Bayer  Akad.  der  Wissen. 
Classe,  Bd.  11,  Abth.  ix,  3-60  and  415-528. 

583.  Waldheim,    Fischer    von.     1840.     Nachtrag    zu 

Herrn  Major  von  Qualen's  geognostischen  Bei- 
tragen  zur  Kenntniss  des  westlicher  Urals.  Bull, 
de  la  Soc.  imp.  des  Natl,  de  Moscou,  488. 

584.  .  1847.  Notice    sur    quelques    Sauriens    de 

l'Oolithe  du  Government  de  Simborsk.  Bull. 
Soc.  Nat.  de  Moscou,  xx,  pt.  1,  364,  pi.  v. 

585.  Walterstorff,  W.  1887.  Ueber    fossile    Froesche 

insbesondere  das  Gattung  Pakeobatrachus.  Theil 
i.Theil  11,  8vd.  Jahresbericht  d.  naturw.  Vereins 
zu  Magdeburg,  1-8 1. 


586.  Walterstorff,  W.  1901.  Ueber  ein  Exemplar  von 

Rana  meriani  von  Meyer  in  Senkenbergischen 
Museum  zu  Frankfurt.  Bericht  Senkenb.  naturf. 
Gesell.,  39-51,  pi. 

587.  Ward,  J.  1890.  On  the  geological  features  of  the 

North  Staffordshire  Coal-Field.  Trans.  North 
Staffordshire  Inst,  of  Mining  and  Mechanical 
Engineers,  x.pl.ix. 

588.  .  1900.  On  the  occurrence  of  Labyrinthodont 

remains  in  the  Keuper  sandstone  of  Stanton. 
Trans.  North  Staffordshire  Field  Club,  xxxiv, 
108-112,  pis.  iv-v. 

589.  .  Paleontology  of  the  Patty  Coal  Field.  Mem. 

Geol  Surv.  United  Kingdom,  pt.  in,  285-357. 

590.  Watson,   D.  M.  S.     1914.    The    Cheirotherium. 

Geol.  Mag.,  vi,  1,  No.  603,  395-398. 

591.  .  1912.  The  larger  Coal  Measure  Amphibia. 

Mem.  and  Proc.  Manchester  Lit.  and  Philos. 
Soc,  lvii,  pt.  1,  No.  1,  1-4,  1  pi. 

592.  — ■ — — .  1913.  Batrachiderpeton  lineatum,  Hancock 

and  Atthey,  a  Coal-Measure  Stegocephalian. 
Proc.  Lond.  Zool.  Soc,  pt.  iv,  949-962,  with  2 
plates  and  figures  in  text. 

593.  .  1913.  The  early  evolution  of  the  Amphibia. 

Rept.  83d  Meeting  British  Assn.  Adv.  Sci.,  532. 

1912.  On  some  Reptilian  lower  jaws.  Ann. 


594- 


595- 


and  Mag.  Nat.  Hist.,  8th  series,  vol.  10,  July-Dec, 
573-587.  with  figures. 

-.  1913.  Micropholis  Stowi  Huxley,  a  temno- 


spondylous  amphibian  from  South  Africa.  Geol. 
Mag.,  5th  Dec,  io,  340-346,  with  figures. 

596.  Weiss,  Ch.  Ernst.     1871.     Paleontologisch  geog- 

nostische  Untersuchungen  aus  dem  Gebirge  auf 
der  Sudseite  der  rheinischen  Devons.  Bonn. 
Sitzungsb.  Niederrhein  Gesell.,  33-35.  (Archego- 
saurus extremity.) 

597.  — — ■ — .  1877.  Ueber  Protriton  petrolei  von  Fried- 

richroda  in  Thuringen.  Zeit.  d.  deutsch.  geol. 
Gesell.,  xxix,  202. 

598.  Wheatley,  Chas.  M.     1871.     The  bone  cave   of 

eastern  Pennsylvania.  Amer.  Jour.  Sc  (3),  I, 
384-385. 

599.  White,  David.  1906.  Geological   position    of   the 

principal  insect-bearing  localities  of  the  American 
Paleozoic,  in  "  Handlirsch's  Revision  of  American 
Paleozoic  Insects. "  Proc.  U.  S.  Nat.  Mus.,  xxix, 
No.  1441,664-667. 

600.  .  1907.  Report  of  the  field  work  in  the  coal 

districts  of  the  state.  Bull.  No.  4,  Illinois  State 
Geol.  Surv.,  201 .  (Gives  correlation  of  the  Mazon 
Creek  Beds.) 

601.  Wiedersheim,  Robert.  1876.  Die  altesten  Formen 

des  Carpus  und  Tarsus  der  heutigen  Amphibien. 
Morphol.  Jahrb.,  Bd.  2, 421-423,  Taf.  xxix,  fig.  2. 

602.  .  1878.   Ueber  Labyrinthodon  ruetemeyeri — 

Anatomie  der  Gehirn.  Abhandl.  d.  schweizer 
paleontol.  Gesell.,  v,  1-56,  mit  3  tafeln. 

-.  1879.  Die  Anatomie  der  Gymnophionen, 


603. 


pp.  i-ioi,  Tafeln  i-ix.  (Refers  to  extinct  Am- 
phibia in  connection  with  phylogeny.) 

604.  .  1886.  Lehrbuch  der  vergleichenden  Anat- 
omie der  Wirbelthiere.    2  Auflage. 

605.  ■  and  W.  N.  Parker.    1897.    Elements  of  the 

comparative  anatomy  of  vertebrates.  Trans- 
lated by  W.  N.  Parker.  Labyrinthodontia,  242, 
fig.  193  c. 

606.  Wilder,  H.  H.  1909.  History  of  the  human  body. 

Amphibia,  541-542. 


BIBLIOGRAPHY  OF  THE   FOSSIL   AMPHIBIA. 


217 


607.  Williston,  S.  W.  1897.  Some    vertebrates    from 

the  Kansas  Permian.  Kans.  Univ.  Quart.,  ser. 
A,  VI,  No.  1,  53,  fig.  Cricotus. 

608.  .  1897.  A    new    Labyrinthodont    from    the 

Kansas  Carboniferous.  Kans.  Univ.  Quart.,  vi, 
No.  4,  ser.  A,  309,  fig.  and  pi.  (Refers  tooth  and 
skull  fragments  to  Mastodonsaurus.) 

1899.    The  Red-Beds  of  Kansas.    Science 


609. 


610. 


611. 


612. 


613. 


614. 


615. 


616. 


617. 


618. 


619. 


620. 


621. 


622. 


(2),  ix,  221. 

.  1899.  Notes    on    the    coraco-scapula    of 

Eryops  Cope.    Kans.  Univ.  Quart.,  vm,  No.  4, 
185-186,  pis.  xxvii-xxx.    (See  Moodie  (476).) 

1900.  Some  fish   teeth  from   the   Kansas 


Cretaceous.  Kans.  Univ.  Quart.,  ser.  A,  ix,  No.  I, 

Jan.,  pi.  vi,  figs.  13,  13a. 
.  1904.  The  temporal  arches  of  the  Reptilia. 

Biol.  Bull.,  vil,  No.  4,  175-192. 
.     1908.    The  faunal  relations  of   the  early 

vertebrates.    Jour.  Geol.,  xvn,  No.  5,  389-402. 

Also,  Outlines  of    geologic    history,  Willis  and 

Salisbury,  1910, 163-175. 
.    1908.    Lysorophus,  a    Permian   Urodele. 

Biol.  Bull.,  xv,  No.  5,  229-240.    Review  in  Rev. 

crit.  de  Paleozoologie  13th  Annee,  No.  2,  82. 
-.  1909.  The  skull  and  extremities  of  Diplo- 


caulus.    Trans.  Kans.  Acad.  Science,  xxii,  122- 
131,  pis.  1-5. 

-.  1909.  New  or  little-known  Permian  verte- 


brates, Trematops,  new  genus.    Jour.  Geol.,  xvn, 
No.  7,  Nov.,  636-658,  7  figs. 

1909.  Principal  characters  of  the  Chelydo- 


sauria,  a  suborder  of  temnospondylous  Amphib- 
ians from  the  Texas  Permian.  A.  A.  A.  S.  Soc. 
Vert.  Paleon. 

.  1910.  Dissorophus.    Jour.  Geol.,  xvm,  No. 

6,  526-535.  Pis-  i-iii- 

.     1910.     Cacops,      Desmospondylus,     new 

genera  of  Permian  Vertebrates.  Bull.  Geol.  Soc. 
Amer.,  xxi,  249-284,  pis.  6-17. 

.  1910.  New  Permian  Reptiles:  Rachitomous 

vertebrae.  Jour.Geol.,xvm,  N0.7, 585-600,  with  figs. 

— — .  191 1.  American  Permian  vertebrates.  Am- 
phibia, 9-14,  pis.  xxxii,  xxxviii. 

1913.    The  primitive  structure  of  the  man- 


623. 
624. 
624a. 


624A. 


6  2  4C. 


dible  in  amphibians  and  reptiles.    Jour.   Geol., 
xxi,  625-627, 1  fig. 

1 9 14.  The  osteology  of  some  American  Per- 


mian vertebrates.  Jour.  Geol.,  xxii,  No.  4,416,  figs. 
1914.  Water  reptiles  of  the  past  and  pres- 


ent.   Amphibia,  35,  47,  and  figures. 

- — ■ .  1914.  Broiliellus,  a  new  genus  of  am- 
phibians from  the  Permian  of  Texas.  Jour.  Geol., 
xxii,  No.  1,  49-56,  figs.  1-2. 

-.    19 1 4.    Restorations   of   some  American 

Permocarboniferous    amphibians    and     reptiles. 
Jour.  Geol.,  xxii,  No.  1,  57-70,  figs.  1-11. 

.  19 1 5.  Trimerorhachis,  a  Permian  temno- 

spondyl  amphibian.     Jour.  Geol.,  xxiii,  No.  3, 
246-255,  figs.  1-6. 
(i2^d.  .  1916.  Synopsis  of  the  Amer.  Permocar- 
boniferous   Tetrapoda.    Contrib.   from    Walker 
Museum,  vol.  I,  No.  9,  pp.  200-211,  figs.  38-53. 

625.  Wiman,  Carl.  1908.  Ein  Paar  Labyrinthodonten- 

reste  aus  der  Trias  Spitzbergens.    Bull.  Geol.  Inst. 
Univ.  of  Upsala,  ix,  34-40,  Tafeln  11,  Nos.  17, 18. 

626. •.  1913.  Ueber  das  Hinterhaupt  der  Laby- 

rinthodonten.     Bull.  Geol.  Inst,  of  Upsala,  XII, 
1-7,  figs.  1-8. 


627. 
628. 

629. 
630. 
631. 

632. 

633- 
°34- 

635- 
636. 

637- 

638. 
639- 

640. 

641. 
642. 

643- 

644- 
645- 


Wiman  Carl.  1914.  LMierdieStegocephalenausder 
Trias  Spitzbergens.  Bull.  Geol.  Inst,  of  Upsala, 
xiii,  1-30,  pis.  i-ix,  figs.  1-10,  bibliography. 

Woodward,  A.  Smith.  1891.  Catalogue  of  the  fos- 
sil fishes  in  the  British  Museum,  pt.  11,  474. 
(Note  on  the  synonymy  of  Pygopterus,  Colosteus, 
and  Archegosaurus. ) 

.    1 89 1.    On  a   Microsaurian   (Hylonomus 

wildii)  from  the  Lancashire  Coal  Field.     Geol. 
Mag.,  n.  s.,  Dec.  in,  vm,  211-213,  with  figs. 

.  1897.  On  a  new  specimen  of  the  Stegoce- 

phalian    Ceraterpeton    galvani    Huxley.     Geol. 
Mag.,  Dec.  iv,  iv,  No.  397,  293,  with  plate. 

1904.  On  two  new  Labyrinthodont  skulls 


of  the  genera  Capitosaurus  and  Aphaneramraa 
(from  the  Trias  of  Spitzbergen).  Proc.  Zool. 
Soc.  London,  170-176,  with  2  pis. 

.  1906.  The    relations    of    paleontology    to 

biology.  Ann.  and  Mag.  Nat.  Hist.,  xvm,  No. 
106,312-318. 

1905.  Fishes  and  Labyrinthodonts.  Paleon- 


tologica  Indica,  n.  s.,  11,  Memoir  No.  2,  10,  2  pis. 
-.  1909.  On  a  new  Labyrinthodont  from  Oil 


Shale,  N.  S.  W.     Rec.  Geol.  Surv.  N.  S.  W.,  vm, 
pt.  iv,  317-319.  pl-  51- 

Woodward,  H.  B.  1887.  The  geology  of  England 
and  Wales.   The  Northumberland  Coal  Field,  1 80. 

Wood  worth,  J.  B.  1900.  Vertebrate  footprints  on 
Carboniferous  shales  of  Plain ville,  Mass.  Bull. 
Geol.  Soc.  Amer.,  xi,  449-454,  pl.  40,  fig.  I. 

Wyman,  Jeffries.  1843.  Analogies  which  exist 
between  the  structure  of  the  teeth  of  the  Lepi- 
dostei  (Gars  and  Gar-pikes)  and  those  of  the  Laby- 
rinthodonts (extinct  Amphibia).  Proc.  Bost. 
Nat.  Hist.  Soc,  1,  131-132;  also  Silliman's  Jour- 
nal, 843. 

.  1853.  Notes    on    the    Reptilian    remains 

(Dendrerpeton).    Quart.  Jour.   Geol.  Soc.  Lon- 
don, ix,  64-66. 

-.  1856.  On   a  batrachian   reptile  from  the 


Coal  Formation.  Proc.  Amer.  Assn.  Adv.  Sci., 
10th  Meeting  at  Albany,  172-173.  Reviews: 
Edinb.  n.  Philos.  Jour.  1857,  v,  360-361;  Neues 
Jahrb.  f.  Mineral.,  Geol.  u.  Paleon.,  1.  857,  340. 
1858.  On  some  remains  of  batrachian  rep- 


tiles discovered  in  the  Coal  Formation  of  Ohio. 
Amer.  Jour.  Science,  xxv  (2),  Mar.,  158-164, 
fig.  Review:  Zeit.  f.  gesammt  Naturwissensch., 
xin,  71 ;  Neues  Jahrb.  f.  Mineral.,  Geol.  u.  Paleon., 
1858,340. 

Yakowlew,  N.  1902.  Neue  Funde  von  Trias — 
Saurier  auf  Spitzbergen.  Verhandl.  Russ.  k.  Min. 
Gesell.,  xl,  180;  Nachtrag,  xli,  165. 

Zittel,  Karl  von.  1 887-1 890.  Handbuch  der 
Paleontologie,  I  Abth.,  Bd.  in,  337~437.  Amphibia. 
Miinchen  u.  Leipsic,  R.  Oldenbourg,  also  East- 
man's translation  of  above,  2,  1 14-139,  figs. 

.    1888.    Ueber  Labyrinthodon  ruetimoycri 

(Wiedersheim).  Neues  Jahrb.  f.  Mineral.,  Geol.u. 
Paleon.,  Bd.  11,  17.  (Regards  the  form  as  a  rep- 
tile.) 

-.  1899.  Amphibien.  Geschichte  der  Geologie 


und  Paleontologie.  Muenchen  u.  Leipzig,  1899. 
827. 
Zwick,  W.  1897.  Beitrage  zur  Kenntniss  des  Baucs 
und  der  Entwickelung  der  Amphibienglie.lmassen, 
besonders  des  Carpus  und  Tarsus.  Zeitschr. 
wissen.  Zool.,  Bd.  63,  62-144,  Taf.  4"5- 


AN  INDEX  TO  THE  BIBLIOGRAPHY  OF  FOSSIL  AMPHIBIA. 


The  following  will  be  of  assistance  to  those  wishing  aid  in  finding  the  literature 
on  special  phases,  whether  anatomical  or  geological,  of  the  fossil  Amphibia. 
The  author's  name  and  the  number  of  his  paper  in  the  preceding  bibliography 
are  given  in  groups  of  classified  subjects,  beginning  with  distribution  and  ending 
with  anatomy.  The  various  anatomical  notes  of  interest  are  especially  widely 
scattered  in  papers  which  often  deal  with  a  variety  of  other  subjects. 


I.  Geological  and  Geographical  Distribution. 

Devonian: 

North  America:  Marsh,  407. 

Europe:  Lohest,  381;  Weiss,  596;  Thevenin,  566. 
Mississippian: 

North  America:  Branson,  50;  Barrell,  21 ;  Lea,  371- 

372- 

Scotland:  Huxley,  331-334;   Atthey,   II;   Hancock 
and  Atthey,  305. 
Coal  Measures  (Pennsylvanian) : 

North  America:  Agassiz,  2;  Butts,  82,  83;  Case,  86, 
94;  Cope,  105,  107,  115,  116,  118,  122,  123,  126, 
127,  131,  145,  152,  167,  176;  Dawson,  200-225; 
Eastman,  230;  Hay,  316;  Jaekel,  347;  King, 
356,  357;  Marsh,  404,  406;  G.  F.  Matthew, 
408-413;  Moodie,  458-465,  469-475,  478,  479, 
482-486;  Mudge,  490;  Newberry,  495-498;  Ray- 
mond, 531;  Schwarz,  540,  541;  Williston,  608; 
Wyman,  639, 640. 

Europe:  Andrews,  8;  Atthey,  11;  Bailey,  12-13; 
Barkas,  15-20;  Bolton,  42-43;  Credner,  179-194 
(see  also  under  Permian);  Davis,  199;  Dunlop, 
229;  Embleton  and  Atthey,  235;  Etheridge, 
241;  Fritsch,  251;  Gaudry,  263-268,  278,  281, 
282;  Gergens,  291;  Goldfuss,  297;  Hancock  and 
Atthey,  304-310;  Lydekker,  394;  von  Meyer, 
422,  426,  436;  Oldham,  500;  Owen,  507-509, 
514,  515;  Schwarz,  540;  Thevenin,  565-568; 
Weiss,  597;  Woodward,  629,  630;  Woodworth, 
636. 

Australia:  Stephens,  557;  Woodward,  634. 
Permian : 

North  America:  Broili,  55,  56,  58,63,65;  Case,  86-93, 
95-100;  Cope,  129,  132-133,  137,  138,  140-144, 
156,  169,  170-175;  Emmons,  237;  Gregory,  299; 
W.  D.  Matthew,  414,  415;  Moodie,  457,  458, 
476,  477;  Sternberg,  558;  Strickler,  564;  Willis- 
ton,  607,  609,  610,  614,  615,  616,  617-624. 

Europe:  Ammon,  7;  Branco,  48;  Broili,  59,  60,  61, 
62,  66;  Credner,  179-195;  Deichmueller,  224, 
225;  Eichwald,  233;  Emery,  236;  Fritsch,  251; 
Gaudry,  258-261,  269-282;  Geinitz,  287;  Geinitz 
und  Deichmueller,  289,  290;  Goldfuss,  295,  296; 
von  Huene,  324-327;  Huxley,  328;  Jaekel,  344, 
345;  LeRoy,  375;  Lloyd,  379;  Lortet,  382;  von 
Meyer,  428,  436,  442;  Roemer,  534;  Traut- 
schold,   573;  Twelvetrees,   576. 

Africa:  Broom,  68-71-73,  74 

Asia:  Lydekker,  384,  386;  Huxley,  333;  Woodward, 
633- 


Triassic : 

North  America:  Branson,  49;  Cope,  106,  ill,  121, 

157;  Emmons,  237;  Leidy,  373;  Lucas,  383. 
Europe:  Alberti,  4;  Burmeister,  80,  81;  Fraas,  242- 

245, 247a;  von  Huene,  323;  Huxley,  329;  Jaeger, 

338-34i;  Metcalfe,  417;  von  Meyer,  421,  425, 

439;  Miall,  453;   Owen,  505;   Quenstedt,   527; 

Seeley,  544;  Storrie,  560;  Ward,  588;  Wieder- 

sheim,  602;  Woodward,  631. 
Spitzbergen:  Wiman,     625-627;     Woodward,     631; 

Yakowlew,  641;  Seeley,  547. 
Africa:  Broom,  69;  Huxley,  330;  Owen,  517,  519-520. 
Asia:  Lydekker,  384-391;  Owen,  510,  511. 
Australia:  Huxley,  330;  Stephens,  554,  555. 

Jurassic: 

Europe:  Dollo,  226. 
Comanchean : 

North  America:  Marsh,  405;  Moodie,  480,  481. 

Europe:  Vidal,  579,  580. 

Cretaceous: 

North  America:  Cope,  128;  Hatcher,  314;  Lambe, 
365,  366;  Williston,  6u. 

Tertiary: 

Europe:  Beyrich,  36;  Bieber,  37;  Cuvier,  197;  Fraas, 
246,  247;  Gunther,  301;  Laube,  367-370;  von 
Meyer,  424,  430,  444,  446,  447;  Portis,  525; 
Scheuchzer,  535;  Tschudi,  574-575;  Walter- 
storff,  585-586. 

Pleistocene: 

North  America:  Brown,  76;  Wheatley,  598. 
South  America:  Lydekker,  393. 
Asia:  Clark,  103;  Lydekker,  393;  Owen,  506. 
Europe:  Lydekker,  393. 


II.    Phvlogenv  of  Amphibia. 

Arldt,  10;  Baur,  22-25,  3'i  Boulenger,  44;  Branson,  49; 
Budgett,  79;  Cope,  153,  166, 173;  Davison, 198;  Dollo, 
228;  Gadow,  256;  Gaudry,  280,  282;  Gregory,  299, 
300,  300a;  Haeckel,  312;  Huxley,  337;  Kingsley,  358; 
Moodie,  4896;  Owen,  512;  Pollard,  524;  Quenstedt, 
527;  Thevenin,  567-568;  Versluys,  578;  Vogt,  581; 
Wiedersheim,  601. 


III.  Ontogeny  of  Amphibia. 

Ammon,  7;  Credner,  187;  Fritsch,  251;  Hay,  315;  von 
Meyer,  422;  Thevenin,  566. 

18 


INDEX  TO   THE    BIBLIOGRAPHY  OF   FOSSIL  AMPHIBIA. 


219 


IV.    Structure  and  Morphology  of  Amphibia. 

I, at  oral-line  System  of  Sensory  Organs:  Andrews,  8;Baur, 
31 ;  Branson,  49;  Fraas,  242,  247a;  Huxley,  333;  Mal- 
branc,  401 ;  Miall,  450;  von  Meyer,  436;  Moodie,  458, 
465,  478,  488;  Thevenin,  568. 

Pineal  Eye:  Cope,  160;  Credner,  187;  Jaekel,  346. 

Morphology  of  the  Skull:  Albrecht,  5;  Baur,  24,  27,  29, 
31;  Boulenger,  44;  Branson,  49;  Broili,  55,  56,  66: 
Case,  85,  93,  98;  Cope,  136,  177;  Credner,  187-191; 
Fraas,  242,  247a;  Fritsch,  251;  Fuchs,  255;  Gaupp, 
283;  Goodrich,  298;  Jaekel,  347;  Maggi,  397.  398: 
von  Meyer,  428,  436,  439;  Moodie,  458,  478;  Owen, 
51 1 ;  Seeley,  546;  Thevenin,  568;  Thyng,  570;  Willis- 
ton,  612,  614,  615,  616,  619;  Wiman,  626;  Wood- 
ward, 631. 

Brain:  Wiedersheim,  602;  Moodie,  487. 

Ear:  Cope,  159;  Broom,  75. 

Eye:  Cope,  105,  107;  Moodie,  478. 

Teeth:  Broili,  58;  Credner,  194;  Fraas,  242;  Fritsch,  251; 
Jaekel,  342;  Owen,  502,  503,  504,  505;  Seeley,  548; 
Strickler,  564;  Tomes,  571-572;  Williston,  608;  Wy- 
man,  637. 

Gills:  Budgett,  79;  Credner,  187;  Cope,  123;  Fritsch, 
251;  Gaudry,  268;  Thevenin,  566-568;  von  Meyer, 
436- 


Vertebrae  and  Ribs:  Baur,  25,  26;  Broili,  61;  Case,  98; 
Cope,  133,  136,  142,  148,  153;  Credner,  i79-<93: 
Fraas,  242;  Fritsch,  251;  Gadow,  256;  Gegenbaur, 
284;  Jaekel,  344,  348;  Lillie,  378:  Marsh,  404:  von 
Meyer,  436;  Mivart,  456;  Moodie,  468;  Schwarz, 
540,  541;  Thevenin,  565-568;  Williston,  620. 

Pectoral  and  Pelvic  Arches:  Broili,  62;  Broom,  72;  Cope, 
161;  Credner,  180-185;  Fritsch,  251;  Gregory,  299, 
3000;  Gegenbaur,  285;  Jaekel,  347;  Moodie,  478; 
Williston,  610,  616. 

Limbs,  Carpus,  and  Tarsus:  Baur,  22,  23,  28;  Cope,  161 ; 
Credner,  180-185;  Emery,  236;  Fritsch,  251;  Greg- 
ory, 299-3000 ;  Jaekel,  347 ;  Wiedersheim,  601 ;  Zwick, 

645- 

Clasping  Organs:  Barkas,  16;  Blanchard,  39;  Braun,  52; 
Hilton,  319;  Moodie,  461;  Fritsch,  251;  Stock,  561; 
Newberry,  498. 

Muscles:  Moodie,  464. 

Alimentary  Canal :  Moodie,  47 1 ,  474,  478. 

Integument,  Scales,  and  Structure  of  Bone:  Jaekel,  343; 
Moodie,  464,  465,  478;  Broili,  62;  Dawson,  208. 

Footprints:  Barrell,  21;  Branson,  50;  Brodie,  54;  Butts, 
82-83;  Dawson,  207,  208;  Fritsch,  251 ;  Geinitz,  288; 
Hickling,  318;  King,  356-357;  Lea,  371,  372;  Leidy, 
374;  Marsh,  406;  G.  F.  Matthew,  408-413;  Owen, 
504;  Moodie,  465;  Pabst,  521;  Smith,  549;  Wood- 
worth,  636. 


INDEX. 


Page. 

Aistopoda Si  77 

Alimentary  Canal 25,  58 

Amblyodon 178 

A.  probleraaticum 179 

American  Museum 1,8 

Ames  Limestone 12 

Amphibamus 127 

A.  grandiceps 2,  7,  15,  128 

A.  thoracatus 132 

Amphibamidae 127 

Amphibia 3 

Classification 46 

Definition 49 

Discovery  in  Carboniferous 6 

Geographic  and  Geologic  Distribution. ...       9 

History  of  Classification 39 

Anaschisma,  Lateral  Line  System  of 35 

Anisodexis 164 

Anthracosauridaa 187 

Apoda 33 

Archegosaurus 1,  7,  35 

Arm 29 

Atlas 27 

Axis 27 

Baphetes 188 

B.  minor 189 

B.  planiceps 6,  188 

Baur,  George 39 

Brachydectes  newberryi 177 

Branchiosauria 4,  50 

Branchiosauridas 51 

Branchiosaurus 53 

Branson,  E.  B 38,  192 

Brown,  N.  H 12 

Budgett,  J.  S 78 

Cannelton  Slates 10,  15 

Carr,  J.  C 13,  14,  22 

Case,  E.  C 9,  182 

Caudata 67 

Cephalerpeton  ventriarmatum 133 

Cercariomorphus  parvisquamis 139 

Clepsydrops  Shales 9 

Cocytinidas 67 

Cocytinus  gyrinoides 68 

Cope,  E.  D 1,7,9,34.42,43,  131,207 

Credner,  Hermann 1 ,  24,  52 

Cricotidae 180 

Ctcnerpeton  alveolatum 168 

Dawson,  Sir  J.  W 7,  8,  19,  20,  32,  66,  194,  197 

Dean,  Bashford 1,  32,  187 

Dendrerpeton I9>  194 

D.  acadianum 194 

D.  oweni 196 


Page. 

Dermal  Appendages 197 

Devonian 10,  37 

Diceratosaurus 8,  1 1 6 

D.  lajvis 121 

D.  punctolineatus 117 

D.  robustus 123 

Diplocaulia 34 

Dromopus 37,  201 

D.  aduncus 37 

D.  agilis 201 

Eobaphetes 191 

E.  kansensis 192 

Eosauravus 1 ,  87,  1 76 

Eosaurus  acadianus 7,  1 89 

Eoserpeton  tenuicorne 1 24 

Erierpeton  branchialis 69 

Erpetobrachium  mazonensis 152 

Erpetosaurus   97 

E.  acutirostris 108 

E.  minutus 105 

E.  obtusus 99 

E.  radiatus 98 

E.  sculptilis 106 

E.  tabulatus 101 

E.  tuberculatus no 

Eryopidae 182 

Eryops 23,  34,  182 

Euamphibia 46,  49 

Eumicrerpeton  parvum 57 

Eurythorax  sublasvis 1 72 

Eye 25,131 

Footprints 15,  201 

Fritsch,  Anaton 5,  54,  77,  78 

Fritschia  curtidentata 85 

Gergens,  Dr 6 

Gurley,  W.  F.  E 9,  12 

Hay,  O.  P 8,  129 

Hussakof,  Louis 1,  16,  172 

Huxley,  Thomas  H 3,  137 

Hylerpeton 83 

H.  dawsonii 83 

H.  intermedium 84 

H.  longidentatum 84 

Hylonomidae 79 

Hylonomus 79 

H.  latidens 81 

H.  lyelli 80 

H.  multidens 81 

H.  wymani 81 

Hylopus  logani 6 

Hyoid 25 

Hyphasma  laevis 70 


220 


INDEX. 


221 


Ichthycantlmte '73 

Ichthycanthus 1 73 

I.  ohiensis 1 73 

I.  platypus 174 

Ichthyerpeton  squamosum 137 

Ja:kel,  Otto 8,  118 

Joggins,  The  South  (Nova  Scotia) 19 

Kammplatten 5 

Kittanning  Coal 15.  l6 

Lacoe,  R.  D.  (Collection  of) 1 

Lateral  Line  System 32 

Leg 30,95 

Lepospondylia 76 

Lcptophractus 169 

L.  dentatus 17l 

L.  lineolatus I71 

L.  obsoletus 169 

Linton,  Ohio,  Coal  Measures 16 

Amphibia  of 18 

Logan,  William 6 

Louisville,  Kansas 10 

Lydckker,  Richard 44,  77 

Lyell,  Charles 19.  20 

Macrerpetida; 183 

Macrerpeton '84 

M.  deani lg6 

M.  huxleyi 184 

Mandible 25,  103 

Marsh,  0.  C 10,  37.  189,  202 

Mastodonsauridx 200 

Mastodonsaurus  sp.  indet 200 

Matthew,  G.  F 8,  22 

Mauch  Chunk 37 

Mazon  Creek  Shales 7,  I2 

Amphibia  of 13 

Mazonerpeton 61 

M.  costatum 63 

M.  longicaudatum 61 

Meyer,  Hermann  von 6 

Micrerpeton 51 

M.  caudatum 52 

Microsauria 4,  34,  7° 

Relation  to  Reptilia 77 

Mississippian  Amphibia 37 

Molgophidae '49 

Molgophis '-W 

M.  brevicostatus '5° 

M.  macrurus '49 

M.  wheatleyi '51 

Morphology  of  Coal  Measures  Amphibia 23 

Muscle 32 

Myocommata 3° 

Nccturus 33.  56 

Newberry,  J.  S.  (Collections  of) 1,  8,  17,  18 

Nyraniids J37 


CEstoccphalus 145 

O.  rectidens 147 

0.  remcx 145 

Operculum 172 

Osage  City,  Kansas 10 

Palate 24,  104 

Parabatrachus 7 

Pectoral  Girdle 29,  107 

Pelion 7,  72 

P.  lyelli 73 

Pcliontida? 72 

Pelvic  Girdle 29 

Phlegethontia 155 

P.  linearis 156 

P.  serpens 156 

Phcenix  Tunnel,  Pennsylvania 10 

Pholidogaster 3 

Pitcairn,  Pennsylvania 12 

Platystegos  loricatum 199 

Pleuroptyx  clavatus 12,  19,  153 

Proteida 67 

Proterpeton  gurleyi 12,  178 

Ptyoniidae H1 

Ptyonius I41 

P.  marshii 143 

P.  nummifer 144 

P.  pectinatus 141 

P.  serrula 144 

P.  vinchellianus 143 


Raniceps. 
Ribs 


7 
27 


Occiput 

Odonterpeton  triangularis . 


25 
in 


Salientia 72 

Saurerpeton  latithorax 165 

Sauropleura '57 

S.  digitata '59 

S.  (Anisodexis)  enchodus 164 

S.  foveata l63 

S.  longidentata l62 

S.  newbenyi l6° 

S.  pauciradiata '6° 

S.  scutellata '58 

Sauropleuridae 3°.  '57 

Scales 31.197 

Schwarz,  Hugo 8>  28 

Sclerotic  plates 24 

Scutes ,62 

Skin l6^,97 

Skull 23 

Smilerpeton  acicdentatum 82 

South  Joggins,  Nova  Scotia 7.  '9 

Sparodus  sp "™ 

Spondylerpeton  spinatum ' 8' 

Stegopida; ' I3 

Stegops "3 

S.  divaricata '  '4 

Stereospondylia 5.  34.  200 

Sternum 3° 


222 


INDEX. 


Tarsus 1/6 

Teeth 25 

Temnospondylia 5,  34,  180 

Thinopus  antiquus 10,  37 

Thyrsidium  fasciculare 147 

Traquair,  R.  H 5 

Triton  walthi 50 

Tuditanidas 86 

Tuditanus 86 

T.  brevirostris 89 

T.  longipes 90 

T.  minimus 92 

T.  punctulatus 87 

T.  walcotti 94 


Twin  Mounds,  Kansas 22 

Udden,  J.  A n 

U  record  yliida? 116 

Ventral  Scutellai 30 

Vertebrae 27,  181 

Vertebral  Column 27 

Wiedersheim,  Robert 30 

Williston,  S.  W 2,  10,  200 

Wyman,  Jeffries 7 

Zamenis  flagellum 5 


THIS  BOOK  IS  DUE  ON  THE  LAST  DATE 
STAMPED  BELOW 


RENEWED  BOOKS  ARE  SUBJECT  TO  IMMEDIATE 
RECALL 


MAR     "  |yyg 


LIBRARY,  UNIVERSITY  OF  CALIFORNIA,  DAVIS 

Book  Slip-50m-12,'64(F772s4)458 


373008 


Moodie,  R.L. 

The  coal  measures 
Amphibia  of  North  America. 


QE867 
M?7 


PHYSICAL 
SCIENCES 
LIBRARY 


LIBRARY 

UNIVERSITY  Of  CALIFORNIA 

DAVIS 


373008 


Moodie,   R.L. 

The  coal  measures 
Amphibia  of  North  America 


Call  Number: 


QE867 
M57 


